Orthoptera: Difference between revisions

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{{Automatic taxobox
{{Automatic taxobox
| taxon = Orthoptera
| taxon = Orthoptera
| fossil_range= [[Carboniferous]]–recent {{fossilrange|359|0}}
| fossil_range= [[Carboniferous]]–recent {{fossilrange|300|0}}
| image = Roesel's bush-cricket (Metrioptera roeselii diluta) male.jpg
| image = Roesel's bush-cricket (Metrioptera roeselii diluta) male.jpg
| image_caption = [[Roesel's bush-cricket]]<br />family [[Tettigoniidae]]
| image_caption = [[Roesel's bush-cricket]]<br />family [[Tettigoniidae]]
Line 54: Line 54:
===Phylogeny===
===Phylogeny===


The Orthoptera are divided into two suborders, [[Caelifera]] and [[Ensifera]], that have been shown to be [[monophyletic]].<ref name="Zhou2010">Zhou Z, Ye H, Huang Y, Shi F. (2010) The phylogeny of Orthoptera inferred from mtDNA and description of ''Elimaea cheni'' (Tettigoniidae: Phaneropterinae) mitogenome. ''J. Genet. Genomics.'' 37(5):315-324</ref><ref name=Gwynne>{{cite journal |last1=Gwynne |first1=Darryl T. |title=Phylogeny of the Ensifera (Orthoptera): a hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care in crickets, katydids, and weta |journal= Journal of Orthoptera Research|date=1995 |volume=4 |issue=4 |pages=203–218 |doi=10.2307/3503478 |jstor=3503478}}</ref><ref>{{cite journal |last1=Flook |first1=P. K. |last2=Rowell |first2=C. H. F. |title=The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences |journal=Molecular Phylogenetics and Evolution |date=1997 |volume=8 |issue=1 |pages=89–103 |doi=10.1006/mpev.1997.0412 |pmid=9242597|bibcode=1997MolPE...8...89F }}</ref> A recent comprehensive phylogeny based on analyses of data from transcriptomes and mitochondrial genomes found the following relationships within Orthoptera.<ref name=Song-2020>{{cite journal |last1=Song |first1=Hojun |last2=Béthoux |first2=Olivier |last3=Shin |first3=Seunggwan  |first4=Alexander |last4=Donath |first5=Harald |last5=Letsch |first6=Shanlin |last6=Liu |first7=Duane D. |last7=McKenna |first8=Guanliang |last8=Meng |first9=Bernhard |last9=Misof |first10=Lars |last10=Podsiadlowski |first11=Xin |last11=Zhou |first12=Benjamin |last12=Wipfler |first13=Sabrina |last13=Simon  |title=Phylogenomic analysis sheds light on the evolutionary pathways towards acoustic communication in Orthoptera. |journal=Nat. Commun. |volume=11 |pages=4939 |year=2020 |doi=10.1038/s41467-020-18739-4 |pmc=7532154 }}</ref>
The Orthoptera are divided into two suborders, [[Caelifera]] and [[Ensifera]], that have been shown to be [[monophyletic]].<ref name="Zhou2010">Zhou Z, Ye H, Huang Y, Shi F. (2010) The phylogeny of Orthoptera inferred from mtDNA and description of ''Elimaea cheni'' (Tettigoniidae: Phaneropterinae) mitogenome. ''J. Genet. Genomics.'' 37(5):315-324</ref><ref name=Gwynne>{{cite journal |last1=Gwynne |first1=Darryl T. |title=Phylogeny of the Ensifera (Orthoptera): a hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care in crickets, katydids, and weta |journal= Journal of Orthoptera Research|date=1995 |volume=4 |issue=4 |pages=203–218 |doi=10.2307/3503478 |jstor=3503478}}</ref><ref>{{cite journal |last1=Flook |first1=P. K. |last2=Rowell |first2=C. H. F. |title=The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences |journal=Molecular Phylogenetics and Evolution |date=1997 |volume=8 |issue=1 |pages=89–103 |doi=10.1006/mpev.1997.0412 |pmid=9242597|bibcode=1997MolPE...8...89F }}</ref> A recent comprehensive phylogeny based on analyses of data from transcriptomes and mitochondrial genomes found the following relationships within Orthoptera.<ref name=Song-2020>{{cite journal |last1=Song |first1=Hojun |last2=Béthoux |first2=Olivier |last3=Shin |first3=Seunggwan  |first4=Alexander |last4=Donath |first5=Harald |last5=Letsch |first6=Shanlin |last6=Liu |first7=Duane D. |last7=McKenna |first8=Guanliang |last8=Meng |first9=Bernhard |last9=Misof |first10=Lars |last10=Podsiadlowski |first11=Xin |last11=Zhou |first12=Benjamin |last12=Wipfler |first13=Sabrina |last13=Simon  |title=Phylogenomic analysis sheds light on the evolutionary pathways towards acoustic communication in Orthoptera. |journal=Nat. Commun. |volume=11 |page=4939 |year=2020 |issue=1 |doi=10.1038/s41467-020-18739-4 |pmid=33009390 |pmc=7532154 |bibcode=2020NatCo..11.4939S }}</ref>


{{clade| style=font-size:90%;line-height:90%
{{clade| style=font-size:90%;line-height:90%
Line 121: Line 121:
[[File:Peruvian Stick Bug.jpg|thumb|[[Proscopiidae|Proscopiidae gen. sp.]] from the [[Andes]] of [[Peru]]]]
[[File:Peruvian Stick Bug.jpg|thumb|[[Proscopiidae|Proscopiidae gen. sp.]] from the [[Andes]] of [[Peru]]]]


Taxonomists classify members of the Caelifera and Ensifera into infraorders and superfamilies as follows:<ref>{{cite web|title=Orthoptera Species File Online|url=https://bugscope.beckman.illinois.edu/pdfs/insects/Orthoptera_Species.pdf |publisher=University of Illinois|access-date=6 January 2018}}</ref><ref name="BlackithBlackith1968">{{cite journal|last1=Blackith|first1= RE|last2=Blackith|first2= RM|title=A numerical taxonomy of Orthopteroid insects|journal=Australian Journal of Zoology|volume=16|issue=1 |year=1968|pages=111 |doi=10.1071/ZO9680111}}</ref><ref name="FlookKlee1999">{{cite journal|last1=Flook|first1=P. K.|last2=Klee|first2=S. |last3=Rowell|first3=C. H. F.|last4=Simon|first4=C.|title=Combined Molecular Phylogenetic Analysis of the Orthoptera (Arthropoda, Insecta) and Implications for Their Higher Systematics|journal=Systematic Biology|volume=48 |issue=2|year=1999 |pages=233–253|issn=1076-836X|doi=10.1080/106351599260274|pmid=12066707|url=http://doc.rero.ch/record/300023/files/48-2-233.pdf|doi-access=free}}</ref><ref>{{Cite web |title=superfamily Gryllotalpoidea Leach, 1815: Orthoptera Species File |url=http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1128272 |access-date=2023-09-01 |website=orthoptera.speciesfile.org}}</ref>   
Taxonomists classify members of the Caelifera and Ensifera into infraorders and superfamilies as follows:<ref>{{cite web|title=Orthoptera Species File Online|url=https://bugscope.beckman.illinois.edu/pdfs/insects/Orthoptera_Species.pdf|publisher=University of Illinois|access-date=6 January 2018|archive-date=2 March 2021|archive-url=https://web.archive.org/web/20210302101804/https://bugscope.beckman.illinois.edu/pdfs/insects/Orthoptera_Species.pdf}}</ref><ref name="BlackithBlackith1968">{{cite journal|last1=Blackith|first1= RE|last2=Blackith|first2= RM|title=A numerical taxonomy of Orthopteroid insects|journal=Australian Journal of Zoology|volume=16|issue=1 |year=1968|page=111 |doi=10.1071/ZO9680111}}</ref><ref name="FlookKlee1999">{{cite journal|last1=Flook|first1=P. K.|last2=Klee|first2=S. |last3=Rowell|first3=C. H. F.|last4=Simon|first4=C.|title=Combined Molecular Phylogenetic Analysis of the Orthoptera (Arthropoda, Insecta) and Implications for Their Higher Systematics|journal=Systematic Biology|volume=48 |issue=2|year=1999 |pages=233–253|issn=1076-836X|doi=10.1080/106351599260274|pmid=12066707|url=http://doc.rero.ch/record/300023/files/48-2-233.pdf|doi-access=free}}</ref><ref>{{Cite web |title=superfamily Gryllotalpoidea Leach, 1815: Orthoptera Species File |url=http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1128272 |access-date=2023-09-01 |website=orthoptera.speciesfile.org}}</ref>   


* Suborder [[Caelifera]] – grasshoppers, pygmy mole crickets and allies
* Suborder [[Caelifera]] – grasshoppers, pygmy mole crickets and allies
Line 153: Line 153:
===As pests===
===As pests===


Several species of Orthoptera are considered pests of crops and rangelands or seeking warmth in homes by humans. The two groups of Orthoptera that cause the most damage are [[grasshopper]]s and [[locust]]s. Locust are historically known for wiping out fields of crops in a day. Locust have the ability to eat up to their own body weight in a single day.<ref name=":0">{{Cite web |url=http://animals.nationalgeographic.com/animals/bugs/locust/ |archive-url=https://web.archive.org/web/20100207174802/http://animals.nationalgeographic.com/animals/bugs/locust/ |url-status=dead |archive-date=February 7, 2010 |title=Locusts, Locust Pictures, Locust Facts - National Geographic |last=Society |first=National Geographic |website=National Geographic |access-date=2016-04-11}}</ref> Individuals gather in large groups called swarms, these swarms can range up to 80 million individuals that stretch 460 square miles.<ref name=":0" /> Grasshoppers can cause major agricultural damage but not to the documented extent as locust historically have. These insects mainly feed on weeds and grasses, however, during times of drought and high population density they will feed on crops. They are a known pest in [[soybean]] fields and will likely feed on these crops once preferred food sources have become scarce.<ref>{{Cite web |url=https://extension.entm.purdue.edu/fieldcropsipm/insects/soybean-grasshopper.php |title=Grasshoppers {{!}} Pests {{!}} Soybean {{!}} Integrated Pest Management {{!}} IPM Field Crops {{!}} Purdue University |last=Krupke |first=Christian  |website=extension.entm.purdue.edu |access-date=2016-04-11}}</ref>
Several species of Orthoptera are considered pests of crops and rangelands or seeking warmth in homes by humans. The two groups of Orthoptera that cause the most damage are [[grasshopper]]s and [[locust]]s. Locusts are historically known for wiping out fields of crops in a day. Locusts have the ability to eat up to their own body weight in a single day.<ref name=":0">{{Cite web |url=http://animals.nationalgeographic.com/animals/bugs/locust/ |archive-url=https://web.archive.org/web/20100207174802/http://animals.nationalgeographic.com/animals/bugs/locust/ |archive-date=February 7, 2010 |title=Locusts, Locust Pictures, Locust Facts - National Geographic |last=Society |first=National Geographic |website=National Geographic |access-date=2016-04-11}}</ref> Individuals gather in large groups called swarms, consisting of up to 80 million individuals and spanning as much as 460 square miles.<ref name=":0" /> Grasshoppers can cause major agricultural damage but not to the documented extent as locusts historically have. These insects mainly feed on weeds and grasses, however, during times of drought and high population density they will feed on crops. They are a known pest in [[soybean]] fields and will likely feed on these crops once preferred food sources have become scarce.<ref>{{Cite web |url=https://extension.entm.purdue.edu/fieldcropsipm/insects/soybean-grasshopper.php |title=Grasshoppers {{!}} Pests {{!}} Soybean {{!}} Integrated Pest Management {{!}} IPM Field Crops {{!}} Purdue University |last=Krupke |first=Christian  |website=extension.entm.purdue.edu |access-date=2016-04-11}}</ref>


===As food===
===As food===
{{See also|Insects as food}}
{{See also|Insects as food}}
Most orthopterans are edible, making up 13% of all insects including some 80 species of grasshoppers being regularly consumed worldwide.<ref name=":1">{{Cite book |last=van Huis |first=Arnold |url=https://www.fao.org/3/i3253e/i3253e.pdf |title=Edible Insects: Future Prospects for Food and Feed Security |isbn=9789251075968 |location=Rome |oclc=868923724 |author-link=Arnold van Huis |pages=13–14}}</ref> In [[Madagascar]] and [[Oaxaca]], grasshoppers and locusts are usually collected early in the morning when it is cooler as the orthopterans are less mobile due to being [[Ectotherm|cold-blooded]].<ref name=":1" /> In Thailand, [[House cricket|house crickets]] are commonly reared and eaten; as of 2012, around 20,000 cricket farmers had farms in 53 of their [[Provinces of Thailand|76 provinces]].<ref name=":1" />
Most orthopterans are edible, making up 13% of all insects including some 80 species of grasshoppers being regularly consumed worldwide.<ref name=":1">{{Cite book |last=van Huis |first=Arnold |url=https://www.fao.org/3/i3253e/i3253e.pdf |title=Edible Insects: Future Prospects for Food and Feed Security |date=2013 |isbn=978-92-5-107596-8 |location=Rome |oclc=868923724 |author-link=Arnold van Huis |pages=13–14}}</ref> In [[Madagascar]] and [[Oaxaca]], grasshoppers and locusts are usually collected early in the morning when it is cooler as the orthopterans are less mobile due to being [[Ectotherm|cold-blooded]].<ref name=":1" /> In Thailand, [[House cricket|house crickets]] are commonly reared and eaten; as of 2012, around 20,000 cricket farmers had farms in 53 of their [[Provinces of Thailand|76 provinces]].<ref name=":1" />


In the second century BCE in [[Ancient Greece]], [[Diodorus Siculus]] is known to have called people from [[Ethiopia]] ''Acridophagi'', meaning "eaters of locusts."<ref name=":1" />
In the second century BCE in [[Ancient Greece]], [[Diodorus Siculus]] is known to have called people from [[Ethiopia]] ''Acridophagi'', meaning "eaters of locusts."<ref name=":1" />


In [[Judaism]], the Orthoptera include the only insects considered [[kosher]]. The list of dietary laws in the [[book of Leviticus]] forbids all flying insects that walk, but makes an exception for [[Kosher locust|certain locusts]].<ref name="Gordon">{{citation |title=The eat-a-bug cookbook |pages=3 |url=https://books.google.com/books?id=sI0Zax6ljq8C&pg=PA3 |first=David George |last=Gordon |publisher=Ten Speed Press |year= 1998 |isbn=978-0-89815-977-6}}</ref> The [[Torah]] states the only kosher flying insects with four walking legs have knees that extend above their feet so that they hop.<ref>{{citation |url= http://bible.ort.org/books/pentd2.asp?action=displayanchor&pentid=P3008 |title=Navigating the Bible: Leviticus }}</ref><!-- Thus nonjumping Orthoptera such as [[mole crickets]] are certainly not kosher.-->
In [[Judaism]], the Orthoptera include the only insects considered [[kosher]]. The list of dietary laws in the [[book of Leviticus]] forbids all flying insects that walk, but makes an exception for [[Kosher locust|certain locusts]].<ref name="Gordon">{{citation |title=The eat-a-bug cookbook |page=3 |url=https://books.google.com/books?id=sI0Zax6ljq8C&pg=PA3 |first=David George |last=Gordon |publisher=Ten Speed Press |year= 1998 |isbn=978-0-89815-977-6}}</ref> The [[Torah]] states the only kosher flying insects with four walking legs have knees that extend above their feet so that they hop.<ref>{{citation |url= http://bible.ort.org/books/pentd2.asp?action=displayanchor&pentid=P3008 |title= Navigating the Bible: Leviticus |access-date= 2010-01-16 |archive-date= 2011-06-10 |archive-url= https://web.archive.org/web/20110610042414/http://bible.ort.org/books/pentd2.asp?action=displayanchor&pentid=P3008 }}</ref><!-- Thus nonjumping Orthoptera such as [[mole crickets]] are certainly not kosher.-->


===As creators of biofuel===
===As creators of biofuel===


With new research showing promise in locating alternative [[biofuel]] sources in the gut of insects, grasshoppers are one species of interest. The insect's ability to break down [[cellulose]] and [[lignin]] without producing [[greenhouse gases]] has aroused scientific interest.<ref>{{Cite journal |last1=Shi |first1=Weibing |last2=Xie |first2=Shangxian |last3=Chen |first3=Xueyan |last4=Sun |first4=Su |last5=Zhou |first5=Xin |last6=Liu |first6=Lantao |last7=Gao |first7=Peng |last8=Kyrpides |first8=Nikos C. |last9=No |first9=En-Gyu |date=January 2013 |title=Comparative Genomic Analysis of the Endosymbionts of Herbivorous Insects Reveals Eco-Environmental Adaptations: Biotechnology Applications |journal=PLOS Genetics |volume=9 |issue=1 |pages=e1003131 |doi=10.1371/journal.pgen.1003131 |pmc=3542064 |pmid=23326236 |doi-access=free }}</ref>
With new research showing promise in locating alternative [[biofuel]] sources in the gut of insects, grasshoppers are one species of interest. The insect's ability to break down [[cellulose]] and [[lignin]] without producing [[greenhouse gases]] has aroused scientific interest.<ref>{{Cite journal |last1=Shi |first1=Weibing |last2=Xie |first2=Shangxian |last3=Chen |first3=Xueyan |last4=Sun |first4=Su |last5=Zhou |first5=Xin |last6=Liu |first6=Lantao |last7=Gao |first7=Peng |last8=Kyrpides |first8=Nikos C. |last9=No |first9=En-Gyu |date=January 2013 |title=Comparative Genomic Analysis of the Endosymbionts of Herbivorous Insects Reveals Eco-Environmental Adaptations: Biotechnology Applications |journal=PLOS Genetics |volume=9 |issue=1 |article-number=e1003131 |doi=10.1371/journal.pgen.1003131 |pmc=3542064 |pmid=23326236 |doi-access=free }}</ref>


==See also==
==See also==

Latest revision as of 23:31, 2 October 2025

Template:Short description Template:Automatic taxobox

Orthoptera (Template:Etymology) is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grasshoppers, locusts, and close relatives; and Ensifera – crickets and close relatives.

More than 20,000 species are distributed worldwide.[1] The insects in the order have incomplete metamorphosis, and produce sound (known as a "stridulation") by rubbing their wings against each other or their legs, the wings or legs containing rows of corrugated bumps. The tympanum, or ear, is located in the front tibia in crickets, mole crickets, and bush crickets or katydids, and on the first abdominal segment in the grasshoppers and locusts.[2] These organisms use vibrations to locate other individuals.

Grasshoppers and other orthopterans are able to fold their wings (i.e. they are members of Neoptera).

Etymology

The name is derived from the Greek Script error: No such module "Lang". Template:Transliteration meaning "straight" and Script error: No such module "Lang". Template:Transliteration meaning "wing".

Characteristics

Orthopterans have a generally cylindrical body, with elongated hindlegs and musculature adapted for jumping. They have mandibulate mouthparts for biting and chewing and large compound eyes, and may or may not have ocelli, depending on the species. The antennae have multiple joints and filiform type, and are of variable length.[2]

The first and third segments on the thorax are larger, while the second segment is much smaller. They have two pairs of wings, which are held overlapping the abdomen at rest. The forewings, or tegmina, are narrower than the hindwings and hardened at the base, while the hindwings are membranous, with straight veins and numerous cross-veins. At rest, the hindwings are held folded fan-like under the forewings. The final two to three segments of the abdomen are reduced, and have single-segmented cerci.[2]

Life cycle

Orthopterans have a paurometabolous lifecycle or incomplete metamorphosis. The use of sound is generally crucial in courtship, and most species have distinct songs.[3] Most grasshoppers lay their eggs in the ground or on vegetation. The eggs hatch and the young nymphs resemble adults, but lack wings and at this stage are often called 'hoppers'. They may often also have a radically different coloration from the adults. Through successive moults, the nymphs develop wings until their final moult into a mature adult with fully developed wings.[2]

The number of moults varies between species; growth is also very variable and may take a few weeks to some months depending on food availability and weather conditions.

Evolution

This order evolved Template:Ma with a division into two suborders – Caelifera and Ensifera – occurring Template:Ma.[4]

Phylogeny

The Orthoptera are divided into two suborders, Caelifera and Ensifera, that have been shown to be monophyletic.[5][6][7] A recent comprehensive phylogeny based on analyses of data from transcriptomes and mitochondrial genomes found the following relationships within Orthoptera.[8]

Template:Clade

Notes

Template:Notelist

Taxonomy

File:Garden locust (Acanthacris ruficornis).jpg
Garden locust (Acanthacris ruficornis), Ghana, family Acrididae
File:Variegated grasshopper (Zonocerus variegatus).jpg
Variegated grasshopper (Zonocerus variegatus), Ghana, family Pyrgomorphidae
File:Peruvian Stick Bug.jpg
Proscopiidae gen. sp. from the Andes of Peru

Taxonomists classify members of the Caelifera and Ensifera into infraorders and superfamilies as follows:[9][10][11][12]

Relationships with humans

As pests

Several species of Orthoptera are considered pests of crops and rangelands or seeking warmth in homes by humans. The two groups of Orthoptera that cause the most damage are grasshoppers and locusts. Locusts are historically known for wiping out fields of crops in a day. Locusts have the ability to eat up to their own body weight in a single day.[13] Individuals gather in large groups called swarms, consisting of up to 80 million individuals and spanning as much as 460 square miles.[13] Grasshoppers can cause major agricultural damage but not to the documented extent as locusts historically have. These insects mainly feed on weeds and grasses, however, during times of drought and high population density they will feed on crops. They are a known pest in soybean fields and will likely feed on these crops once preferred food sources have become scarce.[14]

As food

Script error: No such module "Labelled list hatnote". Most orthopterans are edible, making up 13% of all insects including some 80 species of grasshoppers being regularly consumed worldwide.[15] In Madagascar and Oaxaca, grasshoppers and locusts are usually collected early in the morning when it is cooler as the orthopterans are less mobile due to being cold-blooded.[15] In Thailand, house crickets are commonly reared and eaten; as of 2012, around 20,000 cricket farmers had farms in 53 of their 76 provinces.[15]

In the second century BCE in Ancient Greece, Diodorus Siculus is known to have called people from Ethiopia Acridophagi, meaning "eaters of locusts."[15]

In Judaism, the Orthoptera include the only insects considered kosher. The list of dietary laws in the book of Leviticus forbids all flying insects that walk, but makes an exception for certain locusts.[16] The Torah states the only kosher flying insects with four walking legs have knees that extend above their feet so that they hop.[17]

As creators of biofuel

With new research showing promise in locating alternative biofuel sources in the gut of insects, grasshoppers are one species of interest. The insect's ability to break down cellulose and lignin without producing greenhouse gases has aroused scientific interest.[18]

See also

References

Template:Reflist

External links

Template:Sister project Template:Sister project

Template:Orders of Insects Template:Orthoptera

Template:Taxonbar Template:Authority control

  1. ↑ Script error: No such module "citation/CS1".
  2. ↑ a b c d Script error: No such module "citation/CS1".
  3. ↑ Script error: No such module "citation/CS1".
  4. ↑ Chang H, Qiu Z, Yuan H, Wang X, Li X, Sun H, Guo X, Lu Y, Feng X, Majid M, Huang Y (2020) Evolutionary rates of and selective constraints on the mitochondrial genomes of Orthoptera insects with different wing types. Mol Phylogenet Evol
  5. ↑ Zhou Z, Ye H, Huang Y, Shi F. (2010) The phylogeny of Orthoptera inferred from mtDNA and description of Elimaea cheni (Tettigoniidae: Phaneropterinae) mitogenome. J. Genet. Genomics. 37(5):315-324
  6. ↑ Script error: No such module "Citation/CS1".
  7. ↑ Script error: No such module "Citation/CS1".
  8. ↑ Script error: No such module "Citation/CS1".
  9. ↑ Script error: No such module "citation/CS1".
  10. ↑ Script error: No such module "Citation/CS1".
  11. ↑ Script error: No such module "Citation/CS1".
  12. ↑ Script error: No such module "citation/CS1".
  13. ↑ a b Script error: No such module "citation/CS1".
  14. ↑ Script error: No such module "citation/CS1".
  15. ↑ a b c d Script error: No such module "citation/CS1".
  16. ↑ Script error: No such module "citation/CS1".
  17. ↑ Script error: No such module "citation/CS1".
  18. ↑ Script error: No such module "Citation/CS1".
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