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| binomial_authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
| binomial_authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
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The '''Surinam cockroach''' or '''greenhouse cockroach'''<ref name=gade1997/> ('''''Pycnoscelus surinamensis''''') is a species of burrowing [[cockroach]]. It is a common plant pest endemic to the [[Indomalaya ecozone|Indomalayan]] region that has spread to tropical and into subtropical regions around the world, and in isolated populations to temperate climates where protective habitat such as greenhouses provide shelter for individuals inadvertently shipped in the soil of plants. Its populations are almost exclusively female, and it reproduces asexually, having evolved several clonal strains from its sexual progenitor ''P. indicus''.
The '''Surinam cockroach''' (''Pycnoscelus surinamensis'') is a species of [[cockroach]]. It is approximately 3/4" in length, with uniformly dark brown wings, and a shiny dark brown or black head and pronotum (the shield behind its head).


== Description ==
It is usually not considered a [[pest (animal)|pest]] but it can be found in homes on occasion. It is often brought inside with houseplants. It is found throughout the [[Subtropics |subtropical]] and [[tropical]] climates, including in [[Texas]], [[Louisiana]], and [[Florida]]. As only females are found in the [[United States]], they reproduce by [[parthenogenesis]] there.<ref>D. R. Suiter and P. G. Koehler, [http://edis.ifas.ufl.edu/MG229 Surinam Cockroach, Pycnoscelus surinamensis]</ref><ref>Gary Mullen, Gary Richard Mullen, Lance Durden, [http://books.google.com/books?id=yi6uIQgvFaMC&pg=PA30&lpg=PA30&dq=surinam+cockroach+parthenogenesis&source=bl&ots=7My7htVXXV&sig=pux4fMzyRU6rC8QiSAZGqtTnylI&hl=en&ei=md17Sqxwh-ixA7GFxe8K&sa=X&oi=book_result&ct=result&resnum=10 Medical and Veterinary Entomology]</ref><ref>[http://www.ingentaconnect.com/content/esa/aesa/1967/00000060/00000004/art00010?crawler=true Sexual Isolation in Parthenogenetic Pycnoscelus surinamensis and Application of the Name Pycnoscelus indicus to Its Bisexual Relative (Dictyoptera: Blattaria: Blaberidae: Pycnoscelinae)]</ref>

Adults are around {{convert|18|–|25|mm|in|abbr=on}} in length, and have dark brown to black bodies with shiny brown wings and paler wings.<ref name=mullen2002/><ref name=alford2012>{{cite book|last=Alford|first=David V. |title=Pests of Ornamental Trees, Shrubs and Flowers|url=http://books.google.com/books?id=fWNNjWFwKwgC&pg=PA23|date=30 May 2012|publisher=Manson Publishing|isbn=978-1-84076-628-8|page=23}}</ref> The front edge of the [[pronotum]] (head shield) has a pale white band.<ref name=suiter>{{cite web | last1=Suiter | first1=DR | last2=Koehler | first2=PG | url=http://edis.ifas.ufl.edu/MG229 | title=ENY241/MG229: Surinam Cockroach, Pycnoscelus surinamensis | publisher=Institute of Food and Agricultural Sciences, University of Florida | archiveurl=https://web.archive.org/web/20110825211028/http://edis.ifas.ufl.edu/mg229 | archivedate=25 August 2011 }}</ref> While males are rarely produced, male adults have longer wings than females, completely covering the abdomen,<ref name=schwabe1949/> although both genders are poor flyers.<ref name=bell2007/>

Nymphs are around 4.5 mm long at birth, translucent white with orange-brown mandibles and spines, and darker eye spots than the rest of the head.<ref name=schwabe1949/> The exoskeleton gradually hardens on exposure to air, becoming a glossy brown in 5 to 6 hours, while its underside and legs remain translucent.<ref name=schwabe1949/> Older nymphs develop shiny black anterior abdominal segments, and roughened, dull black posterior segments.<ref name=mullen2002/>

== Range ==

''P. surinamensis'' originated in the Indomalayan region and is now [[Cosmopolitan distribution|cosmopolitan]], found around the world in tropical regions, extending into subtropical regions, and temporarily establishing populations in protective habitats in temperate climates, particularly in [[greenhouse]]s (also called glasshouses) or other areas heated during colder periods.<ref name=mullen2002/><ref name=bell2007>{{ cite book | last1=Bell| first1=WJ | last2=Roth | first2=LM | last3=Nalepa | first3=CA | title=Cockroaches: Ecology, Behavior, and Natural History | year=2007| publisher=JHU Press | isbn=978-0-8018-8616-4 | pages=121–122 | url=http://edu-net.nl/Flora%20en%20fauna/Boeken/Cockroache;%20Ecology,%20behavior%20&%20history%20-%20W.J.%20Bell.pdf | format=PDF }}</ref><ref name=usda1941>{{ cite book | editor=US Department of Agriculture | title=Climate and Man: Part One | url=http://books.google.com/books?id=vPHHk2t1Tq4C&pg=PA524 | edition=Reprinted from the 1941 edition | year=2004 | publisher=University Press of the Pacific | location=Honolulu| isbn=978-1-4102-1538-3 | page=524 }}</ref> It can be spread to greenhouses with shipments of tropical plants.<ref name=usda1941/> In the United States it is common in the Southeastern from North Carolina to Texas, in addition to temporary populations in more temperate climates.<ref name=mullen2002/>

== Habitat ==

''P. surinamensis'' is a burrowing cockroach, commonly burrowing in loose soil, humus, mould, compost piles and lawn thatch, or hiding beneath rocks, rotten branches, trash and other debris.<ref name=mullen2002/><ref name=suiter/><ref name=schwabe1949/><ref name=grandcolas1996/> It is considered peridomestic, found living only near human constructions or crops in a 1996 study, and may be considered [[Synanthrope|synanthropic]].<ref name=grandcolas1996>{{ cite journal | last1=Grandcolas| first1=P.| last2=Dejean | first2=A. | last3=Deleporte | first3=P. | title=The invading parthenogenetic cockroach: a natural history comment on Parker and Niklasson's study | journal=Journal of Evolutionary Biology | volume=9 | issue=6 | year=1996 | pages=1023–1026| issn=1010-061X | doi=10.1046/j.1420-9101.1996.9061023.x | url=http://onlinelibrary.wiley.com/doi/10.1046/j.1420-9101.1996.9061023.x/pdf | format=PDF }}</ref> It has a relatively high rate of cutaneous water loss compared to non-burrowing species of cockroaches, and is nearly exclusively associated with moist soil across its range.<ref name=gade1997>{{cite journal|last1=Gade|first1=B.|last2=Parker | first2=E. D.|title=The effect of life cycle stage and genotype on desiccation tolerance in the colonizing parthenogenetic cockroach Pycnoscelus surinamensis and its sexual ancestor P. indicus|journal=Journal of Evolutionary Biology|volume=10| issue=4|year=1997|pages=479–493|issn=1010-061X| doi=10.1046/j.1420-9101.1997.10040479.x|url=http://onlinelibrary.wiley.com/doi/10.1046/j.1420-9101.1997.10040479.x/pdf}}</ref>

== Pest ==

While the species is occasionally found inside homes, it is not a common household [[pest (animal)|pest]].<ref name=suiter/> It is a plantation and garden pest in tropical regions, and can be especially problematic in heated greenhouses, where large numbers can hide by day and emerge at night, gnawing the softer parts of plants.<ref name=schal1990>{{cite journal | title=Integrated suppression of synanthropic cockroaches | work=Annu. Rev. Entomol. 1990 | volume=35 | pages=521–551 | year=1990 | last1=Schal | first1=C | last2=Hamilton | first2=RL | url=http://www4.ncsu.edu/~coby/schal/1990SchalAREreview.pdf | format=PDF }}</ref><ref name=suiter/> It is thought to be transferred into homes and businesses in potted plants or mulch used for potted plants.<ref name=mullen2002/><ref name=suiter/>

Indoor infestations can often be controlled using sprays or granules on potted plants.<ref name=suiter/> Outdoor treatment should focus on infested locations such as wood piles, mulch, and foundation plantings, and a residual barrier spray may be useful around houses.<ref name=suiter/>

== Parthenogenesis ==

An unusual aspect of ''P. surinamensis'' is that the species reproduces [[parthenogensis|parthenogenetically]], a form of asexual reproduction in which embryos develop from unfertilized eggs. ''P. surinamensis'' reproduce by [[Thelytoky|thelytokous]] parthenogenesis, a type of parthenogenesis in which offspring are almost exclusively female clones. Theylytokous parthenesis is known in about 1% of known animal species.<ref>{{Cite journal| doi = 10.1080/11250008409439455| issn = 0373-4137| volume = 51| issue = 1-2| pages = 1–23| last = White| first = Michael J.D.| title = Chromosomal Mechanisms in Animal Reproduction| journal = Bolletino di zoologia| year = 1984 | url=http://en.wikipedia.org/wiki/Thelytoky | format=PDF}}</ref>

Optional [[Thelytoky|thelkytokous]] parthenogenesis occurs in several cockroach species when females are isolated from males, including in the common domestic pests ''Blatta orientalis'', ''Blattella germanica'', and ''Periplaneta americana'', but in ''P. surinamensis'' it is obligatory parthenogenesis, its sole means of reproduction. Earlier taxonomy treated P. surinamensis as a species with both sexual and asexual forms, reproducing asexually in some populations, but sexually in other populations.<ref name=roth1967>{{ cite journal | title=Sexual Isolation in Parthenogenetic Pycnoscelus surinamensis and Application of the Name Pycnoscelus indicus to Its Bisexual Relative (Dictyoptera: Blattaria: Blaberidae: Pycnoscelinae) | last=Roth | first=Louis M | work=Annals of the Entomological Society of America | volume=60 | issue=4 | date=17 July 1967 | pages=774–779 | publisher=Entomological Society of America | url=http://www.ingentaconnect.com/content/esa/aesa/1967/00000060/00000004/art00010?crawler=true }}</ref><ref name=roth1968/><ref name=mullen2002>{{ cite book | last1=Mullen | first1=Gary R. | last2=Durden | first2=Lance A. |title=Medical and Veterinary Entomology|pages=37,42 | url=http://books.google.com/books?id=yi6uIQgvFaMC&pg=PA37|date=27 September 2002|publisher=Academic Press|isbn=978-0-08-053607-1}}</ref> Roth in 1967 applied the name ''P. indicus'' (Fabricius, 1775) to the sexually-reproducing taxon from which the parthenogenetic form ''P. surinamensis'' is descended.<ref name=roth1968/><ref name=gade1997/> While ''P. surinamensis'' is cosmpolitan, ''P. indicus'' is endemic to the Indomalayasian region and adjacent parts of Southeast Asia, and spread just to [[Hawaii|Hawaiian]] and [[Mauritius|Mauritian]] islands.<ref name=roth1968/>

In ''P. surinamensis'', males are rarely born and are nonfunctional with frequency varying by population. An Australian colony being studied produced several, while none were seen in a Florida colony in over 15 years of study. In a parthenogenetic moth the rate varied by how long a population was stabilized, which suggests the possibility that Australian colony may have developed parthengenesis more recently than the Florida colony.<ref name=roth1968>{{ cite journal| last1=Roth| first1=Louis M.| last2=Cohen| first2=Samuel H.| title=Chromosomes of the Pycnoscelus Indicus and P. Surinamensis Complex (Blattaria: Blaberidae: Pycnoscelinae)| journal=Psyche: A Journal of Entomology| volume=75| issue=1| year=1968| pages=53–76| issn=0033-2615| doi=10.1155/1968/38048 url=http://downloads.hindawi.com/journals/psyche/1968/038048.pdf}}</ref>

P. surinamensis has at least 21 diploid clones, born independently from sexual females, meaning the thelytokous parthenogenesis has evolved repeatedly.<ref name=bell2007/> There are also 11 known triploid clones, produced by backcrosses between clones and ''P. indicus''.<ref name=bell2007/> Different clones have established populations throughout the species' range, with at least ten different clone populations are present in the United States alone.<ref name=roth1968/><ref name=bell2007/>

Obligatory parthenogensis was known only in ''P. surinamensis'' until 2003, when it was reported in a form of the species ''[[Phyllodromica subaptera]]''.<ref name=bell2007/> Asexual forms of ''P. subaptera'', spread through most Mediterranean countries, are morphologically indistinguishable from the sexual form, found only on the Iberian peninsula, suggesting the trait was relatively recently acquired.<ref name=bell2007/>

== Gestation and birth ==

Reproduction is usually [[ovoviviparous]], the ootheca carried internally during gestation until nymphs hatch within the body and are delivered, usually at night or in darkness.<ref name=schwabe1949/> Occasionally stress or other factors may result in premature release of the ootheca, which can result in non-viability of the eggs.<ref name=schwabe1949/> The ootheca itself measures 9x3.5x2.5 mm, housing 30-36 eggs in two rows of alternately spaced segments. Remains of the egg case are expelled around the time the nymphs are being born, sometimes pulled out with with the hind limbs, and the ootheca fragments are usually quickly devoured by the newborn nymphs.<ref name=schwabe1949/>

== Parasite Host ==

''P. surinamensis'' acts as the intermediate host for the parasitic roundworms ''Oxyspirura mansoni'' and ''Oxyspirura parvorum'', which infect the eyes of poultry.<ref name=mullen2002/>

''O. mansoni'', known as the chicken eyeworm among other names,<ref name=schwabe1949>{{ cite journal | last=Schwabe | first=CW | title=Observations on the life history of Pycnoscelus surinamensis (Linn.), the intermediate host of the chicken eyeworm in Hawaii | date=March 1949 | url=http://scholarspace.manoa.hawaii.edu/bitstream/handle/10125/16194/PHES13_433-436.pdf?sequence=1 | format=PDF | work=Proceedings of the Hawaiian Entomological Society | volume=13 | issue=3 | pages=433–436 }}</ref> can cause eye problems ranging from mild conjunctivitis to severe ophthalmia and serious vision impairment in its final hosts, which include chickens, turkeys, guineafowl and peafowl.<ref name=taylor2013>{{ cite book | last1=Taylor | first1=MA | last2=Coop | first2=RL | last3=Wall | first3=RL | title=Veterinary Parasitology | url=http://books.google.com/books?id=tPpEofdsZ5gC&pg=PA1017 | date=13 March 2013 | publisher=John Wiley & Sons | isbn=978-1-118-68711-6 | pages=1017–1018 }}</ref><ref name=usda1941/><ref name=mullen2002/> Its life cycle involves eggs passing through a bird's lachrymal duct, being swallowed and passed in the bird's feces, a ''P. surinamensis'' cockroach eating the feces, larvae emerging in the cockroach, the bird eating the cockroach, and finally the eye worm larvae migrating up the oesophagus and pharynx to the bird's eye.<ref name=taylor2013/> While ''O. mansoni'' is not present in Europe, it is present in many areas of the world, particularly the tropical and subtropical environments where ''P. surinamensis'' is permanently established.<ref name=taylor2013/>


== References ==
== References ==
{{Reflist}}
{{Reflist}}


<!-- ==External links==
<!-- == External Links ==
{{wikispecies|Pycnoscelus surinamensis}}
{{wikispecies|Pycnoscelus surinamensis}}
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{{commons|Pycnoscelus surinamensis}} -->


{{DEFAULTSORT:Surinam Cockroach}}
[[Category:Cockroaches]]
[[Category:Cockroaches]]
[[Category:Animals described in 1758]]
[[Category:Animals described in 1758]]


{{cockroach-stub}}

Revision as of 02:23, 30 December 2013

Surinam cockroach
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
P. surinamensis
Binomial name
Pycnoscelus surinamensis

The Surinam cockroach or greenhouse cockroach[1] (Pycnoscelus surinamensis) is a species of burrowing cockroach. It is a common plant pest endemic to the Indomalayan region that has spread to tropical and into subtropical regions around the world, and in isolated populations to temperate climates where protective habitat such as greenhouses provide shelter for individuals inadvertently shipped in the soil of plants. Its populations are almost exclusively female, and it reproduces asexually, having evolved several clonal strains from its sexual progenitor P. indicus.

Description

Adults are around 18–25 mm (0.71–0.98 in) in length, and have dark brown to black bodies with shiny brown wings and paler wings.[2][3] The front edge of the pronotum (head shield) has a pale white band.[4] While males are rarely produced, male adults have longer wings than females, completely covering the abdomen,[5] although both genders are poor flyers.[6]

Nymphs are around 4.5 mm long at birth, translucent white with orange-brown mandibles and spines, and darker eye spots than the rest of the head.[5] The exoskeleton gradually hardens on exposure to air, becoming a glossy brown in 5 to 6 hours, while its underside and legs remain translucent.[5] Older nymphs develop shiny black anterior abdominal segments, and roughened, dull black posterior segments.[2]

Range

P. surinamensis originated in the Indomalayan region and is now cosmopolitan, found around the world in tropical regions, extending into subtropical regions, and temporarily establishing populations in protective habitats in temperate climates, particularly in greenhouses (also called glasshouses) or other areas heated during colder periods.[2][6][7] It can be spread to greenhouses with shipments of tropical plants.[7] In the United States it is common in the Southeastern from North Carolina to Texas, in addition to temporary populations in more temperate climates.[2]

Habitat

P. surinamensis is a burrowing cockroach, commonly burrowing in loose soil, humus, mould, compost piles and lawn thatch, or hiding beneath rocks, rotten branches, trash and other debris.[2][4][5][8] It is considered peridomestic, found living only near human constructions or crops in a 1996 study, and may be considered synanthropic.[8] It has a relatively high rate of cutaneous water loss compared to non-burrowing species of cockroaches, and is nearly exclusively associated with moist soil across its range.[1]

Pest

While the species is occasionally found inside homes, it is not a common household pest.[4] It is a plantation and garden pest in tropical regions, and can be especially problematic in heated greenhouses, where large numbers can hide by day and emerge at night, gnawing the softer parts of plants.[9][4] It is thought to be transferred into homes and businesses in potted plants or mulch used for potted plants.[2][4]

Indoor infestations can often be controlled using sprays or granules on potted plants.[4] Outdoor treatment should focus on infested locations such as wood piles, mulch, and foundation plantings, and a residual barrier spray may be useful around houses.[4]

Parthenogenesis

An unusual aspect of P. surinamensis is that the species reproduces parthenogenetically, a form of asexual reproduction in which embryos develop from unfertilized eggs. P. surinamensis reproduce by thelytokous parthenogenesis, a type of parthenogenesis in which offspring are almost exclusively female clones. Theylytokous parthenesis is known in about 1% of known animal species.[10]

Optional thelkytokous parthenogenesis occurs in several cockroach species when females are isolated from males, including in the common domestic pests Blatta orientalis, Blattella germanica, and Periplaneta americana, but in P. surinamensis it is obligatory parthenogenesis, its sole means of reproduction. Earlier taxonomy treated P. surinamensis as a species with both sexual and asexual forms, reproducing asexually in some populations, but sexually in other populations.[11][12][2] Roth in 1967 applied the name P. indicus (Fabricius, 1775) to the sexually-reproducing taxon from which the parthenogenetic form P. surinamensis is descended.[12][1] While P. surinamensis is cosmpolitan, P. indicus is endemic to the Indomalayasian region and adjacent parts of Southeast Asia, and spread just to Hawaiian and Mauritian islands.[12]

In P. surinamensis, males are rarely born and are nonfunctional with frequency varying by population. An Australian colony being studied produced several, while none were seen in a Florida colony in over 15 years of study. In a parthenogenetic moth the rate varied by how long a population was stabilized, which suggests the possibility that Australian colony may have developed parthengenesis more recently than the Florida colony.[12]

P. surinamensis has at least 21 diploid clones, born independently from sexual females, meaning the thelytokous parthenogenesis has evolved repeatedly.[6] There are also 11 known triploid clones, produced by backcrosses between clones and P. indicus.[6] Different clones have established populations throughout the species' range, with at least ten different clone populations are present in the United States alone.[12][6]

Obligatory parthenogensis was known only in P. surinamensis until 2003, when it was reported in a form of the species Phyllodromica subaptera.[6] Asexual forms of P. subaptera, spread through most Mediterranean countries, are morphologically indistinguishable from the sexual form, found only on the Iberian peninsula, suggesting the trait was relatively recently acquired.[6]

Gestation and birth

Reproduction is usually ovoviviparous, the ootheca carried internally during gestation until nymphs hatch within the body and are delivered, usually at night or in darkness.[5] Occasionally stress or other factors may result in premature release of the ootheca, which can result in non-viability of the eggs.[5] The ootheca itself measures 9x3.5x2.5 mm, housing 30-36 eggs in two rows of alternately spaced segments. Remains of the egg case are expelled around the time the nymphs are being born, sometimes pulled out with with the hind limbs, and the ootheca fragments are usually quickly devoured by the newborn nymphs.[5]

Parasite Host

P. surinamensis acts as the intermediate host for the parasitic roundworms Oxyspirura mansoni and Oxyspirura parvorum, which infect the eyes of poultry.[2]

O. mansoni, known as the chicken eyeworm among other names,[5] can cause eye problems ranging from mild conjunctivitis to severe ophthalmia and serious vision impairment in its final hosts, which include chickens, turkeys, guineafowl and peafowl.[13][7][2] Its life cycle involves eggs passing through a bird's lachrymal duct, being swallowed and passed in the bird's feces, a P. surinamensis cockroach eating the feces, larvae emerging in the cockroach, the bird eating the cockroach, and finally the eye worm larvae migrating up the oesophagus and pharynx to the bird's eye.[13] While O. mansoni is not present in Europe, it is present in many areas of the world, particularly the tropical and subtropical environments where P. surinamensis is permanently established.[13]

References

  1. ^ a b c Gade, B.; Parker, E. D. (1997). "The effect of life cycle stage and genotype on desiccation tolerance in the colonizing parthenogenetic cockroach Pycnoscelus surinamensis and its sexual ancestor P. indicus". Journal of Evolutionary Biology. 10 (4): 479–493. doi:10.1046/j.1420-9101.1997.10040479.x. ISSN 1010-061X.
  2. ^ a b c d e f g h i Mullen, Gary R.; Durden, Lance A. (27 September 2002). Medical and Veterinary Entomology. Academic Press. pp. 37, 42. ISBN 978-0-08-053607-1.
  3. ^ Alford, David V. (30 May 2012). Pests of Ornamental Trees, Shrubs and Flowers. Manson Publishing. p. 23. ISBN 978-1-84076-628-8.
  4. ^ a b c d e f g Suiter, DR; Koehler, PG. "ENY241/MG229: Surinam Cockroach, Pycnoscelus surinamensis". Institute of Food and Agricultural Sciences, University of Florida. Archived from the original on 25 August 2011.
  5. ^ a b c d e f g h Schwabe, CW (March 1949). "Observations on the life history of Pycnoscelus surinamensis (Linn.), the intermediate host of the chicken eyeworm in Hawaii" (PDF). Proceedings of the Hawaiian Entomological Society. 13 (3): 433–436.
  6. ^ a b c d e f g Bell, WJ; Roth, LM; Nalepa, CA (2007). Cockroaches: Ecology, Behavior, and Natural History (PDF). JHU Press. pp. 121–122. ISBN 978-0-8018-8616-4.
  7. ^ a b c US Department of Agriculture, ed. (2004). Climate and Man: Part One (Reprinted from the 1941 edition ed.). Honolulu: University Press of the Pacific. p. 524. ISBN 978-1-4102-1538-3. {{cite book}}: |edition= has extra text (help)
  8. ^ a b Grandcolas, P.; Dejean, A.; Deleporte, P. (1996). "The invading parthenogenetic cockroach: a natural history comment on Parker and Niklasson's study" (PDF). Journal of Evolutionary Biology. 9 (6): 1023–1026. doi:10.1046/j.1420-9101.1996.9061023.x. ISSN 1010-061X.
  9. ^ Schal, C; Hamilton, RL (1990). "Integrated suppression of synanthropic cockroaches" (PDF). Annu. Rev. Entomol. 1990. 35: 521–551.
  10. ^ White, Michael J.D. (1984). "Chromosomal Mechanisms in Animal Reproduction" (PDF). Bolletino di zoologia. 51 (1–2): 1–23. doi:10.1080/11250008409439455. ISSN 0373-4137.
  11. ^ Roth, Louis M (17 July 1967). "Sexual Isolation in Parthenogenetic Pycnoscelus surinamensis and Application of the Name Pycnoscelus indicus to Its Bisexual Relative (Dictyoptera: Blattaria: Blaberidae: Pycnoscelinae)". Annals of the Entomological Society of America. 60 (4). Entomological Society of America: 774–779.
  12. ^ a b c d e Roth, Louis M.; Cohen, Samuel H. (1968). "Chromosomes of the Pycnoscelus Indicus and P. Surinamensis Complex (Blattaria: Blaberidae: Pycnoscelinae)". Psyche: A Journal of Entomology. 75 (1): 53–76. doi:10.1155/1968/38048 url=http://downloads.hindawi.com/journals/psyche/1968/038048.pdf. ISSN 0033-2615. {{cite journal}}: Check |doi= value (help); Missing pipe in: |doi= (help)
  13. ^ a b c Taylor, MA; Coop, RL; Wall, RL (13 March 2013). Veterinary Parasitology. John Wiley & Sons. pp. 1017–1018. ISBN 978-1-118-68711-6.