Paederus

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Staphyliniformia
Superfamilia: Staphylinoidea

Familia: Staphylinidae
Subfamilia: Paederinae
Tribus: Paederini
Subtribus: Paederina
Genus: Paederus
Subgenera: Anomalopaederus – Eopaederus – Gnathopaederus – Harpopaederus – Heteropaederus – Megalopaederus – Neopaederus – Nepalopaederus – Oreinopaederus – Paederognathus – Poederomorphus – Pseudopaederus

Species overview: P. abdominalis – P. abnormalis – P. adelaidae – P. adiectus – P. adumbratus – P. aequabilis – P. aequilobatus – P. agnatus – P. ahrensi – P. albertisi – P. albipilis – P. albopubescens – P. alesi – P. alfierii – P. algiricus – P. aliiceps – P. almorensis – P. alternans – P. alticola – P. altivagans – P. alutaceus – P. alutipennis – P. alutithorax – P. amazonicus – P. amicus – P. amplicollis – P. andinus – P. anguinus – P. angulicollis – P. angusticeps – P. angustipennis – P. anmamontis – P. annapurnaensis – P. annexus – P. antennocinctus – P. anthracinus – P. antiquus – P. antoniensis – P. apfelsinicus – P. apteromelas – P. aquaticus – P. aquatilis – P. arabicus – P. arduus – P. argentatus – P. argentinus – P. armstrongi – P. arndti – P. arrowi – P. arrowianus – P. atriventris – P. atrocyaneus – P. aureipennis – P. australis – P. baeri – P. bagmatiensis – P. bakeri – P. bakerianus – P. balcanicus – P. balfourbrownei – P. banghaasi – P. basalis – P. basipes – P. basiventris – P. baudii – P. bayeri – P. beccarii – P. benicki – P. bernhauerianus – P. besucheti – P. bhutanicus – P. biacutus – P. bicoloriceps – P. bicoloripes – P. bilobus – P. birmanus – P. bivirgatus – P. blukwensis – P. bohemani – P. bolivianus – P. bonariensis – P. bonus – P. borailicus – P. borneensis – P. brandti – P. brasiliensis – P. braunsi – P. brevicollis – P. brevior – P. brevipennis – P. bruchi – P. brunneolus – P. bucculentus – P. buluensis – P. burgeoni – P. bursavacua – P. busalminensis – P. caffer – P. caligatus – P. cameroni – P. canonicus – P. capensis – P. capitalis – P. caprai – P. captus – P. cariniventris – P. carolinae – P. cartaginis – P. centralis – P. chalybeus – P. cheesmani – P. cheni – P. chimbuensis – P. chinensis – P. chopardi – P. coarctatus – P. coelestinus – P. coerulescens – P. colasi – P. colettae – P. collaris – P. colonus – P. columbinus – P. combustus – P. comorensis – P. compotens – P. condei – P. confusus – P. congoensis – P. conicipennis – P. conicollis – P. conspicuus – P. cooperi – P. copiosus – P. cordovensis – P. coriaceus – P. costaricensis – P. coxalis – P. crassicollis – P. cribricollis – P. cribripennis – P. crinitus – P. cruenticollis – P. cultellatus – P. cumanus – P. currax – P. curticeps – P. curticollis – P. curvatus – P. cyanellus – P. cyaneus – P. cyanocephalus – P. cyanopterus – P. daicongchaoi – P. dammermani – P. darfurensis – P. debilior – P. delkeskampi – P. densiventris – P. deplectens – P. describendus – P. discurrens – P. diversiceps – P. dollmani – P. doriae – P. drakensbergensis – P. drescheri – P. duplex – P. ecuadorensis – P. edentulus – P. edwardsi – P. eidmanni – P. elegans – P. elegantulus – P. elongatus – P. ertli – P. erythraeanus – P. erythroderus – P. excellens – P. excisissimus – P. eximius – P. extraneus – P. fageli – P. fastuosus – P. feae – P. femoralis – P. femoratus – P. ferreirai – P. ferus – P. fijiensis – P. filicornis – P. flaviceps – P. flavitarsis – P. flavocaudatus – P. flavoterminatus – P. floridanus – P. formosanus – P. freudei – P. freyi – P. frontalis – P. fruhstorferi – P. fulgidipennis – P. fulvicornis – P. fulvocaudatus – P. funebris – P. funereus – P. furcillatus – P. furcispinosus – P. fuscipes – P. fuscus – P. gabonicus – P. gebieni – P. gedyei – P. germairei – P. germanus – P. gestroi – P. gigas – P. globulicollis – P. goellnerae – P. gottschei – P. gowdeyi – P. gracilacutus – P. gracilior – P. gracilis – P. grandicollis – P. grandis – P. gratiosus – P. graueri – P. greeni – P. gressitti – P. gridellii – P. haematoderus – P. hagenensis – P. hartmanni – P. hebridensis – P. heynei – P. hiekei – P. himalayicus – P. hingstoni – P. hintzi – P. hlavaci – P. hornabrooki – P. horni – P. huonensis – P. ignotus – P. iheringi – P. ikelaensis – P. illiesi – P. imitator – P. impressipennis – P. incertus – P. incisus – P. inconspicuus – P. incurvatus – P. inexpectatus – P. insularis – P. intermedius – P. iowensis – P. irianensis – P. ismayi – P. itombwensis – P. jacobsoni – P. jaegeri – P. jamaicensis – P. jansei – P. janssensi – P. japonicus – P. jarrigeanus – P. javanus – P. jessoensis – P. jianyueae – P. jilongensis – P. junodi – P. kabarensis – P. kaindiensis – P. kakodanus – P. kambaitiensis – P. kamonoensis – P. kaszabi – P. katrinae – P. katubuensis – P. kilimandjarensis – P. kivuanus – P. kochi – P. koebelei – P. konfuzius – P. kopetzi – P. koreanus – P. kosempoensis – P. kozuensis – P. kraepelini – P. kuborensis – P. kundelungensis – P. kuntzeni – P. kurosawai – P. lacordairei – P. laensis – P. laetipes – P. laetus – P. lamottei – P. langei – P. laoticus – P. lasti – P. lateralis – P. laticeps – P. lativentris – P. lewisi – P. licenti – P. limnophilus – P. lineativentris – P. lineodenticulatus – P. lingualis – P. listeri – P. littoralis – P. littorarius – P. littoreus – P. lividus – P. loebli – P. lombockianus – P. longiceps – P. lootensi – P. luberensis – P. luctuosus – P. luffianus – P. luluensis – P. luridiventris – P. maai – P. macellus – P. machadoanus – P. machadoi – P. madagascariensis – P. madudanus – P. mafuluensis – P. magniceps – P. magnificus – P. magnus – P. malaisei – P. malus – P. mandibularis – P. mangolae – P. marcuzzii – P. margelanicus – P. mariepskopensis – P. marshalli – P. martensi – P. mateui – P. maynei – P. megascutum – P. melampus – P. melanogaster – P. melanopus – P. memnonius – P. mesopotamicus – P. methneri – P. mexicanus – P. meyricki – P. michaelensis – P. micropterus – P. mimicus – P. minicus – P. minimus – P. monstratus – P. montanellus – P. montanus – P. monticola – P. montislomaensis – P. montivagans – P. morio – P. morosus – P. multidenticulatus – P. mussardi – P. mutans – P. mwengensis – P. nakaoi – P. nakurensis – P. nanlingensis – P. natmataungensis – P. neotropicus – P. nepalensis – P. nepalicus – P. niger – P. nigerrimus – P. nigricornis – P. nigricrus – P. nigripennis – P. nigripes – P. nigriventris – P. nigrolineatus – P. noncurvatus – P. nyamukubiensis – P. obliteratus – P. obscuripes – P. ohausi – P. okapaensis – P. opacicollis – P. opacus – P. ornaticornis – P. ovaliceps – P. overlaeti – P. paeninsula – P. pallidulus – P. pallidus – P. palustris – P. papuanus – P. paraguayanus – P. parallelus – P. parcepunctatus – P. parvidenticulatus – P. parvipennis – P. pauloensis – P. pedestris – P. pendleburyi – P. pentagonalis – P. perlongus – P. permixtus – P. perrieri – P. perroti – P. peruanus – P. philippinus – P. piceus – P. pilifer – P. plagiator – P. politus – P. poweri – P. praecellens – P. praeses – P. preangeranus – P. pretiosus – P. problematis – P. protensus – P. pseudobagmatiensis – P. pseudochalybaeus – P. puberulus – P. pubescens – P. pulchellus – P. pumilus – P. punctatus – P. punctiger – P. punctiventris – P. purpuripennis – P. puthzi – P. raffrayi – P. ragusai – P. republicanus – P. reticulatus – P. reticuliventris – P. riftensis – P. riparius – P. roepkei – P. rougemonti – P. ruandanus – P. rubidus – P. rubrothoracicus – P. rudebecki – P. ruficollis – P. rufitarsis – P. rufiventris – P. rufocyaneus – P. rufofasciatus – P. rufotestaceus – P. ruhembeanus – P. rutilicornis – P. ruwenzoricus – P. sabaeus – P. sachtlebeni – P. saegeri – P. saginatus – P. salvini – P. samarensis – P. samoensis – P. samuelsoni – P. santoensis – P. scabripennis – P. schawalleri – P. scheerpeltzi – P. schillhammeri – P. schoenherri – P. schoutedeni – P. schultheissi – P. schulzei – P. schusteri – P. scottianus – P. secretus – P. sedlaceki – P. semaranganus – P. semiflavus – P. semiviridis – P. senegalensis – P. sericeicollis – P. setifer – P. setosus – P. seydeli – P. sharpi – P. signaticornis – P. sijthoffi – P. sikkimensis – P. simillimus – P. simsoni – P. sinisterobliquus – P. sjoestedti – P. smetanai – P. socius – P. solidus – P. solodovnikovi – P. sondaicus – P. sparsus – P. speculifrons – P. spirituslohsei – P. stenopterus – P. straelenianus – P. studti – P. subcoriaceus – P. subnitidus – P. subopacus – P. sudanensis – P. sulawesi – P. sumbaensis – P. superbus – P. szechuanus – P. tambulensis – P. tamulus – P. tandalensis – P. tanjorensis – P. taprobanus – P. tempestivus – P. tenuis – P. testaceitarsis – P. testaceopiceus – P. tibestiensis – P. tibetanus – P. toxopei – P. transvaalensis – P. tricolor – P. tridens – P. trihamatus – P. trimerus – P. tristis – P. tropicus – P. tsaratananus – P. tumidicollis – P. turneris – P. turrialbanus – P. tweedensis – P. ugandensis – P. uganensis – P. uhligorum – P. ulindiensis – P. ullrichi – P. uluguruensis – P. unicolor – P. uruguayensis – P. usagarae – P. usambarae – P. usambaricus – P. ustus – P. variceps – P. variicornis – P. vastus – P. velox – P. versicolor – P. vianai – P. villiersi – P. vinsoni – P. violaceipennis – P. virgifer – P. viridipennis – P. volutobliquus – P. vulcanus – P. vulneratus – P. wadai – P. wagenitzi – P. wagenitzoides – P. wauensis – P. weisei – P. wenzeli – P. willersi – P. wilsoni – P. xanthocerus – P. xinjiangensis – P. xuei – P. xui – P. yaoundensis – P. yuangasanus – P. yucateca – P. yunnanensis – P. zairensis – P. zayasi – P. zhangmuensis – P. zulu

Invalid species names: P. dangchangensis – P. daozhenensis – P. latro – P. melanurus – P. pelikani – P. pseudobaudii – P. symmetricus – P. ussuriensis
Name

Paederus Fabricius, 1775: 268 BHL

Type species: Staphylinus riparius Linnaeus, 1758, fixed by subsequent designation by Latreille, 1810: 427.

Synonymy

Geopaederus Gistel, 1848 [new name for Paederus Fabricius (thought preoccupied by Paederia L. in plants?)]
Type species: Staphylinus ripariusLinnaeus, 1758, fixed by autotypy.
Paederillus Casey, 1905: 62
Type species: Paederus littorarius Gravenhorst, 1806, designated in Blackwelder, 1939: 120.
Leucopaederus Casey, 1905: 67
Type species: Paederus ustus LeConte, 1858, fixed by autotypy.

References
Primary references

Fabricius, J.C. 1775: Systema Entomologiae, sistens insectorum classes, ordines, genera, species... Flensburgh and Leipzig, Library Kortii. Internet Archive BHL

Additional references

Adorno, A. & A. Zanetti, 1999: Description of a new Paederus F., 1775 from Sicily, with systematic, biogeographic and ecological notes on Italian species of the subgenera Paederus s.str. and Eopaederus Scheerpeltz, 1957 (Coleoptera: Staphylinidae). Annales de la Société Entomologique de France (N.S.) 35 (2): 189–201. Reference page.
Assing, V. 2013. Three new species of Paederinae from Northeast India (Coleoptera: Staphylinidae). Linzer biologische Beiträge 45(2): 1561–1570. PDF Reference page.
Assing, V. 2015. On the Harpopaederus fauna of China (Coleoptera: Staphylinidae: Paederinae). Linzer biologische Beiträge 47(1): 163–190. Full article (PDF). Reference page. [Catalogue of the Harpopaederus species of China: p. 164]
Frank, J.H. 1988: Paederus, sensu lato (Coleoptera: Staphylinidae): an index and review of the taxa. Insecta mundi, 2 (2): 97–159. (PDF)
Li, Q-L., Li, L-Z., Gu, F-K. & Peng, Z. 2016. New data on brachypterous Paederus (Coleoptera, Staphylinidae) of mainland China. Zootaxa 4184(3): 576–588. DOI: 10.11646/zootaxa.4184.3.11. Reference page.
Li, X. & Zhou, H.-Z. 2009: A review of Chinese species of the subgenus Paederus s. str. (Coleoptera: Staphylinidae: Paederinae) with description of a new species. Zootaxa, 2083: 46–64. Abstract & excerpt (PDF)
Li, X.; Zhou, H.-Z. 2009b: Phylogenetic analysis and taxonomy of the subgenus Gnathopaederus Chapin and related groups of Paederus s.l. (Coleoptera: Staphylinidae: Paederinae). Invertebrate systematics, 23 (5): 422–444. DOI: 10.1071/IS09015
Peng, Z., Li, Q-L., Li, L-Z. & Gu, F-K. 2015. New data on the subgenus Gnathopaederus of the genus Paederus (Coleoptera: Staphylinidae: Paederinae) of mainland China. Zootaxa 3999(3): 421–429. DOI: 10.11646/zootaxa.3999.3.7. Preview (PDF) Reference page.
Peng, Z., Li, L.Z. & Zhao, M.J. 2015. New data on the subgenus Harpopaederus of the genus Paederus (Coleoptera, Staphylinidae, Paederinae) of mainland China. ZooKeys 495: 143–153. DOI: 10.3897/zookeys.495.9484 Reference page.
Scheerpeltz, O. 1957: Vorläufige Diagnosen einiger neuen paläarktischen Arten und Formen der Gattungen Paederidus Muls. Rey, Paederus Fabr. (mit den neuen Untergattungen Eopaederus, Paederus s. str. nov., Heteropaederus, Dioncopaederus und Oedopaederus), Parameropaederus nov gen., Lobopaederus nov. gen. und Megalopaederus nov. gen. Memorie del Museo di storia naturale della Venezia Tridentina, 11: 447–475.
Li, X.-Y., Zhou, H.-Z. & Solodovnikov, A.Y. 2013. Five new species of the genus Paederus from mainland China, with a review of the Chinese fauna of the subtribe Paederina (Coleoptera: Staphylinidae: Paederinae). Annals of the Entomological Society of America 06(5): 562–574. DOI: 10.1603/AN13008 Reference page.

Paederus is a genus of small beetles of the family Staphylinidae ("rove beetles"). With 622 valid species assigned by 1987 to the subtribe Paederina (Paederus and its close allies), and with all but 148 within Paederus itself,[2] the genus is large. Due to toxins in the hemolymph of some species within this genus, it has given its name to paederus dermatitis, a characteristic skin irritation that occurs if one of the insects is crushed against skin. That name, Paederus dermatitis, is a poor choice because, decades earlier, the affliction had been called dermatitis linearis, a name that works in all languages, not just English, because of its Latin origin; the name Paederus dermatitis is also inappropriate because it has shown to be caused by (a) only a few species of the genus Paederus, but (b) also a few species that belong to closely related genera (that are not Paederus) within the subtribe Paederina.[3] A scholarly paper in 2002 suggested that a Paederus species could have been responsible for some of the ten Plagues of Egypt described in the Bible's Book of Exodus.[4]

Description
Unidentified Paederus spreading its wings, Malaysia

Like other beetles (Coleoptera), rove beetles have hardened forewings that cover the flight wings. At one time, the rove beetle group was known as "Brachyptera" (short wings), because their flight wings are folded under short elytra.[5]

Paederus species are widely distributed around the world.[6] They are much more brightly colored than other rove beetles, with metallic blue- or green-colored elytra and many with bright orange or red on the pronotum and the basal segments of the abdomen. These bright colors may be an example of aposematism, a warning signal to potential predators.[7]

Although most adult rove beetles avoid daylight, Paederus species are active during the day and attracted to bright lights after nightfall.[3]

Paederus eggs are laid singly, in moist habitats. Larvae go through two instars before pupation. Both larvae and adults are predatory on other insects.[3] Because of their preference for moist soil, large numbers of Paederus beetles may be attracted to irrigated farmland, where they provide some benefit by eating herbivorous insects but can cause problems for people working in fields or grassy areas.[7]
Pederin
Main article: Pederin

According to Medical and Veterinary Entomology (2009) "At least 20 of the more than 600 species of Paederus beetles have been associated with Paederus dermatitis",[7] even though Paederus beetles do not bite or sting.[8] This skin irritation results from contact with pederin, a vesicant toxin in the hemolymph of many but not all females in the genus Paederus.[9] The toxin is manufactured, not by the beetles themselves, but by endosymbiont bacteria, probably some species of Pseudomonas.[10]
Impact on human beings
Paederus littoralis, Portugal
Main article: Paederus dermatitis

Paederus dermatitis is caused when a pederin-containing beetle is crushed, even partially, against the skin.[7] This skin irritation is also called "dermatitis linearis" or "linear dermatitis" because one can inadvertently drag a beetle across the skin in a more-or-less straight line when trying to brush it away. The resulting inflammation will also be linear.[11] Because Paederus species are widely dispersed around the world, this syndrome has many different local nicknames including "whiplash dermatitis", "spider lick",[7] and "Nairobi fly dermatitis". In East Africa, conjunctivitis from getting pederin in the eye is called "Nairobi eye".[12]

Once pederin is on the skin from the initial beetle contact, it may also be spread elsewhere on the skin. "Kissing" or "mirror-image" lesions where two skin areas come in contact (for example, the elbow flexure) are often seen.[11] Washing the hands and skin with soap and water is strongly recommended if contact with a rove beetle has occurred.[8]

Initial skin contact with pederin shows no immediate result. Within 12–36 hours, however, a reddish rash (erythema) appears, which develops into blisters. Irritation, including crusting and scaling, may last from two to three weeks.[8]
Mass infestations

An article in The Lancet suggests that events like those described as the first two of the ten plagues of Egypt (anoxic die-off in the Nile, followed by many dead frogs) would have created ideal breeding conditions for P. alfierii. The authors suggest that the plague of "boils" could be the skin irritation, typically delayed by a day or more, resulting from contact with Paederus during the third or fourth plagues (lice or flies). They also note that Paederus infestations are often localized, so it would be quite possible for them to have invaded Pharaoh's palace but not the homes of the Jews.[4]

Many modern occurrences of localized but intense Paederus impact are documented in research papers with titles like "An outbreak of paederus dermatitis in a suburban hospital in Sri Lanka",[13] "An outbreak of 268 cases of Paederus dermatitis in a toy-building factory in central China",[14] "Outbreak of dermatitis linearis caused by Paederus ilsae and Paederus iliensis (Coleoptera: Staphylinidae) at a military base in Iraq",[15] and "Evacuation of an Aboriginal community in response to an outbreak of blistering dermatitis induced by a beetle (Paederus australis)".[16]
References

"Paederus Fabricius 1775". Fauna Europaea. Retrieved 28 July 2011.
Frank, J.H. (1988). "Paederus, sensu lato (Coleoptera: Staphylinidae): An index and review of the taxa". Insecta Mundi. 2 (2): 97–159.
Frank, JH; Kanamitsu, K (1987). "Paederus, sensu lato (Coleoptera: Staphylinidae): natural history and medical importance". Journal of Medical Entomology. 24 (2): 155–91. doi:10.1093/jmedent/24.2.155. PMID 3295241.
Norton, Scott A; Lyons, Christina (2002). "Blister beetles and the ten plagues". The Lancet. 359 (9321): 1950. doi:10.1016/S0140-6736(02)08759-7. PMID 12057588. S2CID 38364604.
"7.4 Blister beetles, species". Institute of Tropical Medicine. Archived from the original on 26 September 2012. Retrieved 30 July 2011. "Among the Staphylinidae, the genus Paederus is known to contain at least 25 species."
Sharp, David (1887). Biologia Centrali-Americana. Insecta. Coleoptera Vol. 1 Part 2. Vol. 1. p. 609.
Mullen, Gary; Gary Richard Mullen; Lance Durden (2009). Medical and Veterinary Entomology. Academic Press. p. 102. ISBN 978-0-12-372500-4. Retrieved 31 July 2011. "Pederin contacts human skin only when a beetle is brushed vigorously over the skin or crushed."
"Just the facts…Paederus Beetles" (PDF). US Army Public Health Command. Archived from the original (PDF) on 16 March 2012. Retrieved 30 July 2011.
Kellner, Rupert L. L.; Dettner, Konrat (1996). "Differential efficacy of toxic pederin in deterring potential arthropod predators of Paederus (Coleoptera: Staphylinidae) offspring". Oecologia. 107 (3): 293–300. Bibcode:1996Oecol.107..293K. doi:10.1007/BF00328445. PMID 28307257. S2CID 44776265. "This study investigates the effects of pederin, a hemolymph toxin that is accumulated in the eggs of most Paederus females"
Piel, Jörn (2002). "A polyketide synthase-peptide synthetase gene cluster from an uncultured bacterial symbiont of Paederus beetles". Proceedings of the National Academy of Sciences. 99 (22): 14002–7. Bibcode:2002PNAS...9914002P. doi:10.1073/pnas.222481399. PMC 137826. PMID 12381784.
Singh, Gurcharan; Ali, Syed Yousuf (2007). "Paederus dermatitis". Indian Journal of Dermatology, Venereology and Leprology. 73 (1): 13–5. doi:10.4103/0378-6323.30644. PMID 17314440.
Rapini, Ronald P.; Bolognia, Jean L.; Jorizzo, Joseph L. (2007). Dermatology: 2-Volume Set. St. Louis: Mosby. p. 1180. ISBN 978-1-4160-2999-1.
Kamaladasa, Satheeka D.; Perera, W.D.H.; Weeratunge, L. (1997). "An outbreak of paederus dermatitis in a suburban hospital in Sri Lanka". International Journal of Dermatology. 36 (1): 34–6. doi:10.1046/j.1365-4362.1997.00009.x. PMID 9071612. S2CID 39540236.
Huang, Changzheng; Liu, Yeqiang; Yang, Jing; Tian, Jin; Yang, Lingyun; Zhang, Jing; Li, Yanqiu; Li, Jiawen; et al. (2009). "An outbreak of 268 cases of Paederus dermatitis in a toy-building factory in central China". International Journal of Dermatology. 48 (2): 128–31. doi:10.1111/j.1365-4632.2009.03876.x. PMID 19200185. S2CID 12349799.
Davidson, SA; Norton, SA; Carder, MC; Debboun, M (2009). "Outbreak of dermatitis linearis caused by Paederus ilsae and Paederus iliensis (Coleoptera: Staphylinidae) at a military base in Iraq". U.S. Army Medical Department Journal: 6–15. PMID 20084733.
Todd, RE; Guthridge, SL; Montgomery, BL (1996). "Evacuation of an Aboriginal community in response to an outbreak of blistering dermatitis induced by a beetle (Paederus australis)". The Medical Journal of Australia. 164 (4): 238–40. doi:10.5694/j.1326-5377.1996.tb94150.x. PMID 8604198. S2CID 40346362.