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{{Wikispecies|Coccinella novemnotata}}
{{Wikispecies|Coccinella novemnotata}}


'''''Coccinella novemnotata''''', the '''nine-spotted ladybug''' or '''nine-spotted lady beetle''' or '''C9''', is a species of [[Coccinellidae|ladybug]] in the family [[Coccinellidae]], native to North America. Historically, this beetle was once ubiquitous across the continent but experienced a sharp and drastic decline around the 1960’s<ref>{{Cite journal |last=Losey |first=John E. |last2=Perlman |first2=Jordan E. |last3=Hoebeke |first3=E. Richard |date=2007-12-01 |title=Citizen scientist rediscovers rare nine-spotted lady beetle, Coccinella novemnotata, in eastern North America |url=https://doi.org/10.1007/s10841-007-9077-6 |journal=Journal of Insect Conservation |language=en |volume=11 |issue=4 |pages=415–417 |doi=10.1007/s10841-007-9077-6 |issn=1572-9753}}</ref>. Now, considered a rare species, the nine-spotted ladybug has received much attention from researchers who wish to understand the causes of its decline and restore its population to utilize their [[Biological pest control|aphidophagous]] nature, characteristic of the predominantly predatory [[Coccinellidae]], as [[Biological pest control|biocontrol agents]] in agriculture<ref name=":0">{{Cite journal |last=Seago |first=Ainsley E. |last2=Giorgi |first2=Jose Adriano |last3=Li |first3=Jiahui |last4=Ślipiński |first4=Adam |date=2011-07-01 |title=Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data |url=https://www.sciencedirect.com/science/article/pii/S1055790311001540 |journal=Molecular Phylogenetics and Evolution |volume=60 |issue=1 |pages=137–151 |doi=10.1016/j.ympev.2011.03.015 |issn=1055-7903}}</ref>.
'''''Coccinella novemnotata''''', the '''nine-spotted ladybug''' or '''nine-spotted lady beetle''', is a species of [[Coccinellidae|ladybug]] native to [[North America]].


== Taxonomy ==
Although this family has been the focus of many studies in the 19th and 20th century to delineate the subfamily and tribe relations, there is no current consensus as most studies based on morphology contradict. However, a few recent studies based on [[Sequencing|molecular sequencing]] data have allowed for more detailed intense phylogenetic trees to be produced<ref name=":0" />.''Coccinella novemnotata'' belongs to the subfamily [[Coccinellinae]] and genus Coccinella. This family falls under the superfamily [[Cucujoidea]] and suborder [[Polyphaga]]. A number of recent [[Molecular phylogenetics|molecular phylogeny]] studies agree with pre-existing morphology-based relationships that [[Coccinellidae]] are a monophyletic group within Cucujiformia in the Cerylonid series, sister to a clade in [[Endomychidae]] or [[Corylophidae]]. Further molecular phylogentics studies are needed to delineate and establish subfamiliy level trees. Two recent studies present evidence that [[Coccinellinae]] is the only monophyletic subfamily, opposed to [[Coccidulinae]], [[Epilachninae]], [[Scymninae]] and [[Chilocorinae]] which are paraphyletic groups. In addition, there was evidence the tribe Chilocorini is sister to [[Coccinellinae]]<ref>{{Cite journal |last=Magro |first=A. |last2=Lecompte |first2=E. |last3=Magné |first3=F. |last4=Hemptinne |first4=J. -L. |last5=Crouau-Roy |first5=B. |date=2010-03-01 |title=Phylogeny of ladybirds (Coleoptera: Coccinellidae): Are the subfamilies monophyletic? |url=https://www.sciencedirect.com/science/article/pii/S1055790309004163 |journal=Molecular Phylogenetics and Evolution |volume=54 |issue=3 |pages=833–848 |doi=10.1016/j.ympev.2009.10.022 |issn=1055-7903}}</ref>.


==Identification==
==Description and Identification==
The nine-spotted ladybug can be identified by the presence of four black spots on each of its [[elytra]], a single spot split between the elytra, and a black suture between the elytra. Its [[pronotum]] is black, with two connected white marks at the front of its head.
The nine-spotted ladybug can be identified by the presence of four black spots on each of its [[elytra]], a single spot split between the elytra, and a black suture between the elytra. Its [[pronotum]] is black, with two connected white marks at the front of its head. They range from 4.7-7.0 millimeters (0.19-0.28 inches) long and the [[elytra]] can range from yellow to orange and spotted or spotless<ref>{{Cite book |last=Gordon |first=R.D. |title=The Coccinellidae (Coleoptera) of America North of Mexico |publisher=New York Entomological Society |year=1985}}</ref><ref name=":1">{{Cite journal |last=Dobzhansky |first=T. |date=1930 |title=The North American beetles of the genus Coccinella |url=https://doi.org/10.5479/si.00963801.80-2904. |journal=Proceedings of The United States National Museum |volume=80 |pages=1-32}}</ref>. Males can be distinguished by a spot on their anterior [[Arthropod leg|coxae]] and stripe on the [[Arthropod leg|femora]]<ref name=":1" />.


== Life Cycle and Behavior ==
==Range==
''Coccinella novemnotata'' is a [[bivoltine]] species whose [[Diapause|diapauses]] are dependent mostly on photoperiod length and less on temperature and prey availability. Adults enter diapause through the summer and winter and emerge in the autumn and early spring, respectively. They reproduce in early spring and oviposit in early autumn. Adult females determine the [[Ovipositor|oviposition]] sites and eggs are laid in clutches attached to thin branches, leaves, or other surrounding solid material<ref name=":1" />. C9 undergo a [[Holometabolism|holometabolous]] life cycle, beginning from an egg, larvae (4 instars), prepupa, pupa, and adult. ''C. novemnotata'' and other beetles in the subfamily [[Coccinellinae]] lack a covering during the pupal stage, instead, the larval skin is simply peeled off till the cauda. The two most significant sources of mortality are [[Intraguild predation|interspecific predation]] and [[Intraguild predation|cannibalism]]. Larvae are documented to predate on smaller, earlier instar larvae. Eggs, pupae, and molting individuals are significantly more vulnerable, in this sessile state to hunting beetles<ref>{{Cite journal |last=S. Hesler |first=Louis |last2=McNickle |first2=Ginger |last3=A. Catangui |first3=Michael |last4=E. Losey |first4=John |last5=A. Beckendorf |first5=Eric |last6=Stellwag |first6=Leonard |last7=M. Brandt |first7=Danielle |last8=B. Bartlett |first8=Pamela |date=2012-09-07 |title=Method for Continuously Rearing Coccinella Lady Beetles (Coleoptera: Coccinellidae) |url=https://benthamopen.com/ABSTRACT/TOENTOJ-6-42 |journal=The Open Entomology Journal |volume=6 |issue=1 |doi=10.2174/1874407901206010042}}</ref>. In times of general prey scarcity, [[intraguild predation]] is shown to increase as eggs are better sources of nutrients crucial for larval development compared to aphids. Some studies show evidence that ''C. novemnotata'' suffers from higher rates of egg predation by other species like the ''[[Coccinella septempunctata]]'' and [[Intraguild predation|cannibalism]] of its own eggs<ref>{{Cite journal |last=Turnipseed |first=Rakim |last2=Ugine |first2=Todd A. |last3=Losey |first3=John E. |date=2015-06-19 |title=Egg Predation by the Introduced Lady Beetle, Coccinella septempunctata (Coleoptera: Coccinellidae), Lowers Mortality but Raises Relative Risk for the Native Lady Beetle, Coccinella novemnotata |url=https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0118493 |journal=PLOS ONE |language=en |volume=10 |issue=6 |pages=e0118493 |doi=10.1371/journal.pone.0118493 |issn=1932-6203 |pmc=PMC4474596 |pmid=26090935}}</ref>.
''C. novemnotata'' has historically been native in North America to the United States and southern Canada.<ref name="Losey2007"/> ''C. novemnotata'' has become rare across its native range. It was once the most commonly collected coccinellid in the northeastern United States until the early 1990s with the last individuals collected for Maryland, Pennsylvania, and Delaware between 1986 and 1988, and another collection in Maine in 1992.<ref name="Harmon">{{cite journal|author=Harmon, J.P.|display-authors=et al.|year=2007|title=The decline of native coccinellids (Coleoptera: Coccinellidae) in the United States and Canada|journal=Journal of Insect Conservation|volume=11|pages=85–94|doi=10.1007/s10841-006-9021-1}}</ref> Another specimen would not be collected in northeastern states until 14 years later in 2006.<ref name="Losey2007">{{cite journal|author=Losey, J.E.|display-authors=et al.|year=2007|title=Citizen scientist rediscovers rare nine-spotted lady beetle, Coccinella novemnotata, in eastern North America|journal=Journal of Insect Conservation|volume=11|pages=415–417|doi=10.1007/s10841-007-9077-6}}</ref><ref name="Losey2011">{{cite journal|author=Losey, J.E.|display-authors=et al.|year=2007|title=Lady Beetles in New York: Insidious Invasions, Erstwhile Extirpations, and Recent Rediscoveries|journal=Northeastern Naturalist|volume=21|pages=271–284|doi=10.1656/045.021.0209}}</ref> More recently, ''C. novemnotata'' has only been collected sporadically in the Midwest and west coast of the United States.<ref name="Harmon"/>


==Decline==
== Habitat ==
The nine-spotted ladybeetle are nomadic and can thrive in a great variety of available habitats across the continent. Based on several factors such as aphid density, adult distribution, and seasonal availability of herbaceous material to facilitate breeding<ref>{{Cite book |last=Hodek |first=I. |title=Ecology of coccinellidae |last2=Honěk |first2=A. |publisher=Springer Dordrecht |year=2013 |isbn=978-94-017-1349-8 |edition=1 |doi=10.1007/978-94-017-1349-8}}</ref>. Within agricultural landscapes, ''Coccinella novemnotata'' has been reported in cotton, alfalfa, corn, and soybean as a candidate for [[Biological pest control|biological control]] of aphids. In addition, they can be found in suburban parks and gardens. In more natural settings, they are found in open areas with shrubbery and small trees ([[deciduous]] or [[Conifer|coniferous]]), meadows, prairie grassland, and [[Riparian zone|riparian zones]] <ref name=":2">{{Cite web |last=Canada |first=Environment and Climate Change |date=2016-12-09 |title=COSEWIC Assessment and Status Report on the Nine-spotted Lady Beetle Coccinella novemnotata Canada - 2016 |url=https://www.canada.ca/en/environment-climate-change/services/species-risk-public-registry/cosewic-assessments-status-reports/nine-spotted-lady-beetle-2016.html |access-date=2023-11-14 |website=www.canada.ca}}</ref>.
Invasive coccinellid species such as ''[[Harmonia axyridis]]'' are sometimes associated with the decline of ''C. novemnotata'', but the species was becoming rare in some areas before the introduction and population increase of invasive lady beetles. Changing agricultural habitats have also been considered as another possible explanation, but recent studies do not suggest a correlation between ''C. novemnotata'' densities and land-use change.<ref name="Harmon"/>

==Historic and Current Distribution==
''C. novemnotata'' has historically been native in North America to the United States and southern Canada.<ref name="Losey2007" /> In the 1970s-1980’s, ''Coccinella novemnotata'' was reported to be one of the most prevalent species in the [[Northeastern United States|Northeastern]] and continental US, and the southern regions of Canada. In Canada, the nine-spotted ladybeetle was historically abundant in the southern regions spanning from [[Vancouver Island]] to [[Quebec]]. One paper from 1998 reports sightings in [[Guatemala]] and [[Mexico]]<ref name=":3">{{Cite web |last=Almquist |first=D. T. |date=July 11, 2011 |title=Coccinella novemnotata Nine-spotted Lady Beetle |url=https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.113893/Coccinella_novemnotata |access-date=2023-11-14 |website=explorer.natureserve.org}}</ref>.

''C. novemnotata'' has become rare across its native range. It was once the most commonly collected coccinellid in the [[Northeastern United States|Northeastern]] United States until the early 1990s with the last individuals collected for Maryland, Pennsylvania, and Delaware between 1986 and 1988, and another collection in Maine in 1992. A significant effort of [[citizen science]] originating from [[Cornell University]] in New York has reported tens of sightings of the rare beetle in western states of the US, with nearly no sighting on the eastern side<ref name=":3" />. In 2006, a singular C9 was located in [[Washington, D.C.|Washington, DC]], after 14 years of disappearance. In 2008, over 40 individual ''C. novemnotata'' and collected over 30 live specimens, exclusively from the western states<ref>{{Cite journal |last=Losey |first=John |last2=Perlman |first2=Jordan |last3=Kopco |first3=James |last4=Ramsey |first4=Samuel |last5=Hesler |first5=Louis |last6=Evans |first6=Edward |last7=Allee |first7=Leslie |last8=Smyth |first8=Rebecca |date=2012-04 |title=Potential causes and consequences of decreased body size in field populations of Coccinella novemnotata |url=https://linkinghub.elsevier.com/retrieve/pii/S1049964411003355 |journal=Biological Control |language=en |volume=61 |issue=1 |pages=98–103 |doi=10.1016/j.biocontrol.2011.12.009}}</ref>. Another specimen would not be collected in [[Northeastern United States|Northeastern]] states until 14 years later in 2006<ref name="Losey2007">{{cite journal|author=Losey, J.E.|display-authors=et al.|year=2007|title=Citizen scientist rediscovers rare nine-spotted lady beetle, Coccinella novemnotata, in eastern North America|journal=Journal of Insect Conservation|volume=11|pages=415–417|doi=10.1007/s10841-007-9077-6}}</ref><ref name="Losey2011">{{cite journal|author=Losey, J.E.|display-authors=et al.|year=2007|title=Lady Beetles in New York: Insidious Invasions, Erstwhile Extirpations, and Recent Rediscoveries|journal=Northeastern Naturalist|volume=21|pages=271–284|doi=10.1656/045.021.0209}}</ref>. More recently, ''C. novemnotata'' has only been collected sporadically in the [[Midwestern United States|Midwest]] and west coast of the United States.<ref name="Harmon">{{cite journal |author=Harmon, J.P. |display-authors=et al. |year=2007 |title=The decline of native coccinellids (Coleoptera: Coccinellidae) in the United States and Canada |journal=Journal of Insect Conservation |volume=11 |pages=85–94 |doi=10.1007/s10841-006-9021-1}}</ref> A 2015 Canadian report shows larger populations in provinces of British Colombia and Alberta, and smaller sightings in [[Ontario]]<ref name=":3" />.

==Conservation Status==
Invasive coccinellid species and changing agricultural habitats have been considered as possible explanations to their drastic decline, but recent studies do not suggest a correlation between ''C. novemnotata'' densities and invasive species density and land-use change.<ref name="Harmon" />

''Coccinella novemnotata'' is not protected by the U.S. [[Endangered Species Act of 1969|Endangered Species Act]]. Currently, [[NatureServe]] bestows a G5 global rank on ''Coccinella novemnotata'' as a secure species at a low possibility of extinction. The Committee on the Status of Endangered Wildlife in Canada ([[Committee on the Status of Endangered Wildlife in Canada|COSEWIC]]) and Canadian Species at Risk Act (SARA) listed the nine-spotted lady bug as endangered in 2016 and 2023, respectively. Canada [[NatureServe]] ranks the beetle as N2 or imperiled<ref name=":2" />.

While there are numerous hypothesized contributors to their stark decline, no singular cause has been identified so far. The introduction and establishment of the nonnative ''[[Harmonia axyridis]]'' (HA) and ''[[Coccinella septempunctata]]'' (C7) has come to dominate up to 90% of the present coccinellid population in some areas. A general 71% increase in nonnative species was documented in 2013, which not only compete for resources but may be a significant source of [[intraguild predation]]. However, numerous studies provide evidence that the boom in invasive species only slightly coincides with their decline. Pesticides like [[Neonicotinoid|neonicotinoids]] and [[pyrethroid]]-based regulators have synergistic negative effects on lady beetles if they come in contact or consume these substances through prey or plant material<ref name=":3" /><ref name=":2" />. Other than pesticides, habitat loss through abandoned agricultural plots and further [[urbanization]] could further hinder the successful reproduction and survival of these ladybeetles.

While efforts to document the nine-spotted ladybug are extensive compared to other declining species, their mobile and generalist nature hinder concerted efforts of conservation<ref name=":2" />. Although they are only present in low abundance and as a rare member compared to other species within the aphidophagous body of coccinellids, it is hypothesized that the local and minute differences in dietary or environmental niches of other, more popular species like C7 or HA allow for a small population of ''Coccinella novemnotata'' to coexist. Land heterogeneity, even the introduction and establishment of non-native weeds capable of supporting aphids, combined with their generalism may contribute to their continued presence. Although evidently less abundant, populations of ''Coccinella novemnotata'' may be more stable despite their significant decline<ref>{{Cite journal |last=Evans |first=Edward W. |date=2017-12-05 |title=Fates of Rare Species under Siege from Invasion: Persistence of Coccinella novemnotata Herbst in Western North America alongside an Invasive Congener |url=http://journal.frontiersin.org/article/10.3389/fevo.2017.00152/full |journal=Frontiers in Ecology and Evolution |volume=5 |doi=10.3389/fevo.2017.00152 |issn=2296-701X}}</ref>.


==State insect of New York==
==State insect of New York==

Revision as of 11:10, 14 November 2023

Coccinella novemnotata
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Infraorder: Cucujiformia
Family: Coccinellidae
Genus: Coccinella
Species:
C. novemnotata
Binomial name
Coccinella novemnotata
Herbst, 1793

Coccinella novemnotata, the nine-spotted ladybug or nine-spotted lady beetle or C9, is a species of ladybug in the family Coccinellidae, native to North America. Historically, this beetle was once ubiquitous across the continent but experienced a sharp and drastic decline around the 1960’s[1]. Now, considered a rare species, the nine-spotted ladybug has received much attention from researchers who wish to understand the causes of its decline and restore its population to utilize their aphidophagous nature, characteristic of the predominantly predatory Coccinellidae, as biocontrol agents in agriculture[2].

Taxonomy

Although this family has been the focus of many studies in the 19th and 20th century to delineate the subfamily and tribe relations, there is no current consensus as most studies based on morphology contradict. However, a few recent studies based on molecular sequencing data have allowed for more detailed intense phylogenetic trees to be produced[2].Coccinella novemnotata belongs to the subfamily Coccinellinae and genus Coccinella. This family falls under the superfamily Cucujoidea and suborder Polyphaga. A number of recent molecular phylogeny studies agree with pre-existing morphology-based relationships that Coccinellidae are a monophyletic group within Cucujiformia in the Cerylonid series, sister to a clade in Endomychidae or Corylophidae. Further molecular phylogentics studies are needed to delineate and establish subfamiliy level trees. Two recent studies present evidence that Coccinellinae is the only monophyletic subfamily, opposed to Coccidulinae, Epilachninae, Scymninae and Chilocorinae which are paraphyletic groups. In addition, there was evidence the tribe Chilocorini is sister to Coccinellinae[3].

Description and Identification

The nine-spotted ladybug can be identified by the presence of four black spots on each of its elytra, a single spot split between the elytra, and a black suture between the elytra. Its pronotum is black, with two connected white marks at the front of its head. They range from 4.7-7.0 millimeters (0.19-0.28 inches) long and the elytra can range from yellow to orange and spotted or spotless[4][5]. Males can be distinguished by a spot on their anterior coxae and stripe on the femora[5].

Life Cycle and Behavior

Coccinella novemnotata is a bivoltine species whose diapauses are dependent mostly on photoperiod length and less on temperature and prey availability. Adults enter diapause through the summer and winter and emerge in the autumn and early spring, respectively. They reproduce in early spring and oviposit in early autumn. Adult females determine the oviposition sites and eggs are laid in clutches attached to thin branches, leaves, or other surrounding solid material[5]. C9 undergo a holometabolous life cycle, beginning from an egg, larvae (4 instars), prepupa, pupa, and adult. C. novemnotata and other beetles in the subfamily Coccinellinae lack a covering during the pupal stage, instead, the larval skin is simply peeled off till the cauda. The two most significant sources of mortality are interspecific predation and cannibalism. Larvae are documented to predate on smaller, earlier instar larvae. Eggs, pupae, and molting individuals are significantly more vulnerable, in this sessile state to hunting beetles[6]. In times of general prey scarcity, intraguild predation is shown to increase as eggs are better sources of nutrients crucial for larval development compared to aphids. Some studies show evidence that C. novemnotata suffers from higher rates of egg predation by other species like the Coccinella septempunctata and cannibalism of its own eggs[7].

Habitat

The nine-spotted ladybeetle are nomadic and can thrive in a great variety of available habitats across the continent. Based on several factors such as aphid density, adult distribution, and seasonal availability of herbaceous material to facilitate breeding[8]. Within agricultural landscapes, Coccinella novemnotata has been reported in cotton, alfalfa, corn, and soybean as a candidate for biological control of aphids. In addition, they can be found in suburban parks and gardens. In more natural settings, they are found in open areas with shrubbery and small trees (deciduous or coniferous), meadows, prairie grassland, and riparian zones [9].

Historic and Current Distribution

C. novemnotata has historically been native in North America to the United States and southern Canada.[10] In the 1970s-1980’s, Coccinella novemnotata was reported to be one of the most prevalent species in the Northeastern and continental US, and the southern regions of Canada. In Canada, the nine-spotted ladybeetle was historically abundant in the southern regions spanning from Vancouver Island to Quebec. One paper from 1998 reports sightings in Guatemala and Mexico[11].

C. novemnotata has become rare across its native range. It was once the most commonly collected coccinellid in the Northeastern United States until the early 1990s with the last individuals collected for Maryland, Pennsylvania, and Delaware between 1986 and 1988, and another collection in Maine in 1992. A significant effort of citizen science originating from Cornell University in New York has reported tens of sightings of the rare beetle in western states of the US, with nearly no sighting on the eastern side[11]. In 2006, a singular C9 was located in Washington, DC, after 14 years of disappearance. In 2008, over 40 individual C. novemnotata and collected over 30 live specimens, exclusively from the western states[12]. Another specimen would not be collected in Northeastern states until 14 years later in 2006[10][13]. More recently, C. novemnotata has only been collected sporadically in the Midwest and west coast of the United States.[14] A 2015 Canadian report shows larger populations in provinces of British Colombia and Alberta, and smaller sightings in Ontario[11].

Conservation Status

Invasive coccinellid species and changing agricultural habitats have been considered as possible explanations to their drastic decline, but recent studies do not suggest a correlation between C. novemnotata densities and invasive species density and land-use change.[14]

Coccinella novemnotata is not protected by the U.S. Endangered Species Act. Currently, NatureServe bestows a G5 global rank on Coccinella novemnotata as a secure species at a low possibility of extinction. The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) and Canadian Species at Risk Act (SARA) listed the nine-spotted lady bug as endangered in 2016 and 2023, respectively. Canada NatureServe ranks the beetle as N2 or imperiled[9].

While there are numerous hypothesized contributors to their stark decline, no singular cause has been identified so far. The introduction and establishment of the nonnative Harmonia axyridis (HA) and Coccinella septempunctata (C7) has come to dominate up to 90% of the present coccinellid population in some areas. A general 71% increase in nonnative species was documented in 2013, which not only compete for resources but may be a significant source of intraguild predation. However, numerous studies provide evidence that the boom in invasive species only slightly coincides with their decline. Pesticides like neonicotinoids and pyrethroid-based regulators have synergistic negative effects on lady beetles if they come in contact or consume these substances through prey or plant material[11][9]. Other than pesticides, habitat loss through abandoned agricultural plots and further urbanization could further hinder the successful reproduction and survival of these ladybeetles.

While efforts to document the nine-spotted ladybug are extensive compared to other declining species, their mobile and generalist nature hinder concerted efforts of conservation[9]. Although they are only present in low abundance and as a rare member compared to other species within the aphidophagous body of coccinellids, it is hypothesized that the local and minute differences in dietary or environmental niches of other, more popular species like C7 or HA allow for a small population of Coccinella novemnotata to coexist. Land heterogeneity, even the introduction and establishment of non-native weeds capable of supporting aphids, combined with their generalism may contribute to their continued presence. Although evidently less abundant, populations of Coccinella novemnotata may be more stable despite their significant decline[15].

State insect of New York

The nine-spotted ladybug has been the state insect of New York since 1989,[16] though its numbers have declined as the numbers of introduced species such as the seven-spotted ladybug and Asian lady beetle have increased. It was for some time thought extinct in New York,[17] so in 2006, the state considered designating the pink spotted ladybug as state insect instead, but the bill did not pass the Senate. In 2006 the nine-spotted ladybug was rediscovered in Virginia (the first East Coast sighting in 14 years).[18] In 2011, about 20 of these ladybugs were found on a farm in Amagansett, New York, the first such sighting in the state since 1982.[19]

Notes

  1. ^ Losey, John E.; Perlman, Jordan E.; Hoebeke, E. Richard (2007-12-01). "Citizen scientist rediscovers rare nine-spotted lady beetle, Coccinella novemnotata, in eastern North America". Journal of Insect Conservation. 11 (4): 415–417. doi:10.1007/s10841-007-9077-6. ISSN 1572-9753.
  2. ^ a b Seago, Ainsley E.; Giorgi, Jose Adriano; Li, Jiahui; Ślipiński, Adam (2011-07-01). "Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data". Molecular Phylogenetics and Evolution. 60 (1): 137–151. doi:10.1016/j.ympev.2011.03.015. ISSN 1055-7903.
  3. ^ Magro, A.; Lecompte, E.; Magné, F.; Hemptinne, J. -L.; Crouau-Roy, B. (2010-03-01). "Phylogeny of ladybirds (Coleoptera: Coccinellidae): Are the subfamilies monophyletic?". Molecular Phylogenetics and Evolution. 54 (3): 833–848. doi:10.1016/j.ympev.2009.10.022. ISSN 1055-7903.
  4. ^ Gordon, R.D. (1985). The Coccinellidae (Coleoptera) of America North of Mexico. New York Entomological Society.
  5. ^ a b c Dobzhansky, T. (1930). "The North American beetles of the genus Coccinella". Proceedings of The United States National Museum. 80: 1–32.
  6. ^ S. Hesler, Louis; McNickle, Ginger; A. Catangui, Michael; E. Losey, John; A. Beckendorf, Eric; Stellwag, Leonard; M. Brandt, Danielle; B. Bartlett, Pamela (2012-09-07). "Method for Continuously Rearing Coccinella Lady Beetles (Coleoptera: Coccinellidae)". The Open Entomology Journal. 6 (1). doi:10.2174/1874407901206010042.
  7. ^ Turnipseed, Rakim; Ugine, Todd A.; Losey, John E. (2015-06-19). "Egg Predation by the Introduced Lady Beetle, Coccinella septempunctata (Coleoptera: Coccinellidae), Lowers Mortality but Raises Relative Risk for the Native Lady Beetle, Coccinella novemnotata". PLOS ONE. 10 (6): e0118493. doi:10.1371/journal.pone.0118493. ISSN 1932-6203. PMC 4474596. PMID 26090935.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
  8. ^ Hodek, I.; Honěk, A. (2013). Ecology of coccinellidae (1 ed.). Springer Dordrecht. doi:10.1007/978-94-017-1349-8. ISBN 978-94-017-1349-8.
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  13. ^ Losey, J.E.; et al. (2007). "Lady Beetles in New York: Insidious Invasions, Erstwhile Extirpations, and Recent Rediscoveries". Northeastern Naturalist. 21: 271–284. doi:10.1656/045.021.0209.
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  15. ^ Evans, Edward W. (2017-12-05). "Fates of Rare Species under Siege from Invasion: Persistence of Coccinella novemnotata Herbst in Western North America alongside an Invasive Congener". Frontiers in Ecology and Evolution. 5. doi:10.3389/fevo.2017.00152. ISSN 2296-701X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
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