Bipalium

Bipalium is a genus of large predatory land planarians. They are often loosely called "hammerhead worms" or "broadhead planarians" because of the distinctive shape of their head region. Land planarians are unique in that they possess a "creeping sole" on their ventral side.[2] Several species are considered as invasive to the United States[3] and to Europe.[4][5] Some studies have begun the investigation of the evolutionary ecology of these invasive planarians.[6]

Bipalium
Model of Bipalium strubelli Graff, 1899
Scientific classification
Kingdom:
Animalia
Phylum:
Platyhelminthes
Class:
Rhabditophora
Order:
Tricladida
Suborder:
Continenticola
Family:
Geoplanidae
Subfamily:
Bipaliinae
Genus:
Bipalium

Type species
Bipalium fuscatum
Stimpson, 1857
Synonyms
  • Sphyrocephalus Schmarda, 1859
  • Dunlopea Wright, 1860
  • Bipaliura Stimpson, 1861 (misspelling?)
  • Perocephalus von Graff, 1896 (in part)
  • Placocephalus von Graff, 1896 (in part)

Description

The genus Bipalium was initially defined by Stimpson to include land planarians with the head broadened, forming a head plate. Later, in 1899, Ludwig von Graff divided it into three genera according to the shape of the head:[7]

  1. Bipalium: with a well-developed head plate, much broader than long, and with elongated lateral auricles
  2. Perocephalus: rudimentary head plate, not much broader than the body
  3. Placocephalus: flat head plate with a circular outline

Josef Müller, in 1902, considered that no sufficient anatomical basis existed for this distinction, and reunited them under Bipalium. Later, von Graff accepted Müller's conclusions.[7]

Towards the end of the 20th century, Robert E. Ogren and Masaharu Kawakatsu started a series of publications called "The Land Planarian Index series" in which they reviewed and organized all taxonomic information regarding land planarians. At first, they retained all broad-headed planarians in the genus Bipalium, but later split them into four genera based on the anatomy of the reproductive organs: Bipalium, Novibipalium, Humbertium, and Diversibipalium.[8]

Under this scenario, the genus Bipalium is defined as containing broad-headed land planarians with simple copulatory organs, without accessory ducts or copulatory bursa, and with a fold of tissue separating the male and female exit ducts.[8]

Etymology

The name Bipalium comes from Latin bi-, “two” + pala, “shovel” or “spade”, because species in this genus resemble a pickaxe.[9]

Species

The genus Bipalium currently includes the following species:

  • Bipalium adensameri Graff, 1899
  • Bipalium admarginatum de Beauchamp, 1933
  • Bipalium adventitium Hyman, 1943
  • Bipalium alternans de Beauchamp, 1930
  • Bipalium bergendali (Graff, 1899)
  • Bipalium cantori (Wright, 1860)
  • Bipalium choristosperma de Beauchamp, 1925
  • Bipalium crassatrium de Beauchamp, 1939
  • Bipalium distinguendum Müller, 1907
  • Bipalium dubium Loman, 1890
  • Bipalium ephippium Loman, 1890
  • Bipalium fuscatum Stimpson, 1857
  • Bipalium fuscolineatum Kaburaki, 1922
  • Bipalium gestroi Graff, 1894
  • Bipalium glandiantrum Kawakatsu, Sluys & Ogren, 2005
  • Bipalium glaucum (Kaburaki, 1922)
  • Bipalium gracile Loman, 1890
  • Bipalium graffi Müller, 1903
  • Bipalium haberlandti Graff, 1899
  • Bipalium hilgendorfi (Graff, 1899)
  • Bipalium interruptum Graff, 1899
  • Bipalium javanum Loman, 1883
  • Bipalium kaburakii Kawakatsu, Sluys & Ogren, 2005
  • Bipalium katoi Kawakatsu, Sluys & Ogren, 2005
  • Bipalium kewense Moseley, 1878
  • Bipalium kisoense Kaburaki, 1922
  • Bipalium koreense Frieb, 1923
  • Bipalium kraepelini (Ritter-Záhony, 1905)
  • Bipalium marginatum Loman, 1887
  • Bipalium mjobergi de Beauchamp, 1925
  • Bipalium monolineatum Kaburaki, 1922
  • Bipalium moseleyi Loman, 1887
  • Bipalium muninense Kawakatsu, Sluys & Ogren, 2005
  • Bipalium myadenosum de Beauchamp, 1939
  • Bipalium nigrum (Ritter-Záhony, 1905)
  • Bipalium nobile Kawakatsu & Makino, 1982
  • Bipalium ochroleucum Kaburaki, 1922
  • Bipalium pennsylvanicum Ogren, 1987
  • Bipalium penrissenicum Kawakatsu, Ogren & Froehlich, 1998
  • Bipalium penzigi Müller, 1903
  • Bipalium persephone de Beauchamp, 1939
  • Bipalium poiense de Beauchamp, 1925
  • Bipalium rigaudi Graff, 1894
  • Bipalium robiginosum Graff, 1899
  • Bipalium semperi (Graff, 1899)
  • Bipalium simrothi Loman, 1895
  • Bipalium strubelli Graff, 1899
  • Bipalium sudzuki Kawakatsu, 1986
  • Bipalium tetsuyai Kawakatsu, Sluys & Ogren, 2005
  • Bipalium univittatum Grube, 1866
  • Bipalium vagum Jones & Sterrer, 2005
  • Bipalium virile Müller, 1903
  • Bipalium weismanni Ritter-Záhony, 1905
  • Bipalium wiesneri Graff, 1899

Feeding habits

Bipalium species are predatory. Some species prey on earthworms, while others may also feed on mollusks.[10][11] These flatworms can track their prey.[12] When captured, earthworms begin to react to the attack, but a flatworm uses the muscles in its body, as well as sticky secretions, to attach itself to the earthworm to prevent escape. The planarians cover, or cap, the prostomium, peristomium, and anterior end to stop the violent reaction by the earthworm.[13]

To feed on their prey, species of Bipalium often evert their pharynges from their mouths, located on the midventral portion of their bodies, and secrete enzymes that begin the digestion of the prey. The liquefied tissues are sucked into the branching gut of the flatworms by ciliary action.[14]

Reproduction

Reproduction in Bipalium may be asexual or sexual and all species are hermaphroditic.

B. adventitium reproduces sexually and creates egg capsules, which hatch around 3 weeks post-deposition. The egg capsules have a tough exterior and generally contain multiple juveniles.[6]

B. kewense have rarely been observed using egg capsules as a primary method of reproduction. Asexual fragmentation is its main reproductive strategy in temperate regions and most individuals never develop sexual organs.[15] Juveniles of this species, unlike B. adventitium, do not appear the same coloration as parents in their early days.[16]

Toxicity

Recently, the presence of tetrodotoxin, a very potent neurotoxin, was recorded in B. adventitium and B. kewense.[17] It is the first record of tetrodotoxin in terrestrial invertebrates.

Invasive species
Drawings of the anterior region of 6 species of the genus Bipalium, 5 of which are invasive species, showing the differences in shape and color.

Little is known about the ecology of terrestrial planarians, but research has been done on different genera and species, including several native and invasive species in Brazil,[18] Arthurdendyus triangulatus,[19] Rhynchodemus [20] and Bipalium.[21]

Currently, four invasive species of Bipalium are known in the United States: B. adventitium, B. kewense, B. pennsylvanicum, and B. vagum.[11] These planarians are thought to have come to the US on horticultural plants.

Bipalium kewense has been found commonly in American greenhouses since 1901.[22] This species is a voracious predator of earthworms, and has been identified as a nuisance in the southern USA in earthworm-rearing beds.[23] Control of the species is difficult due to the lack of predators. As noted by the University of Florida IFAS department,

Other animals rarely devour land planarians, since surface secretions appear distasteful, if not toxic. Protozoans, including flagellates, ciliates, sporozoans, and nematodes have been detected in land planarians. Because of their cannibalistic habit, land planarians may be their own worst enemy.[24]

Specimens of Bipalium adventitium are characterized by a single dark dorsal stripe. They were first discovered in the US in California and New York[25] but have been found in Illinois[26] as well as most northern states[6] Bipalium kewense have five dark dorsal stripes and a partial dark collar. They have undergone several name changes since their discovery in North America. B. kewense is commonly found across the southern regions of the US.[16] B. kewense is found worldwide in tropical and subtropical regions.[15] Bipalium pennsylvanicum is characterized by its dark brown head and three dorsal stripes. Thus far, it has only been found in Pennsylvania and in coastal South Carolina in 2014.[27] Bipalium vagum is characterized by two dark dorsal blotches on the head, a thick black band around the neck, and three dark dorsal stripes. This species has recently been found in several Gulf Coast states.[11] Recently it was found in Mumbai, India.

  • Bipalium kewense sighted in Austin, Texas
  • Close up of head of Bipalium kewense
  • Specimen of Bipalium found in Georgia, United States
  • Hammerhead worm found in Kerala, India
  • Hammerhead worm found in Chennai, India
  • A hammerhead worm seen in Kolkata, India
gollark: ~~or for misrepresenting eggs you have~~
gollark: Also, four random kind of messy and weird random eggs.
gollark: Expect:100G silverrandom hatchlingsCB gold taken down while you're staring at it in confusion
gollark: Nobody actually *reads* that.
gollark: ~~gets banned for putting text in want box~~

References
  1. Stimpson (1857). "Prodromus descriptionis animalium evertebratorum quæ in Expeditione ad Oceanum, Pacificum Septentrionalem a Republica Federata missa, Johanne Rodgers Duce, observavit er descripsit. Pars I. Turbellaria Dendrocœla". Proc. Acad. Nat. Sci. Phila. 1857: 25.
  2. Curtis, S.K.; Cowden, R.R.; Moore, J.D.; Robertson, J.L. (1983). "Histochemical and ultrastructural features of the epidermis of land planarian Bipalium adventitium". Journal of Morphology. 175 (2): 171–194. doi:10.1002/jmor.1051750206. PMID 30060639.
  3. Ogren, R.E. 1985. The human factor in the spread of an exotic land planarian in Pennsylvania. Proc. of the Penn. Acad. of Sci. 59: 117-118.
  4. Justine, Jean-Lou; Winsor, Leigh; Gey, Delphine; Gros, Pierre; Thévenot, Jessica (2018). "Giant worms chez moi! Hammerhead flatworms (Platyhelminthes, Geoplanidae, Bipalium spp., Diversibipalium spp.) in metropolitan France and overseas French territories". PeerJ. 6: e4672. doi:10.7717/peerj.4672. ISSN 2167-8359. PMC 5969052. PMID 29844951.
  5. Filella-Subira, E (1983). ""Nota sobre la presència de la planària terrestre Bipalium kewense Moseley, 1878 a Catalunya"". Butll. Inst. Cat. Hist. Nat. 49: 151.
  6. Ducey, P. K.; West, L. J.; Shaw, G.; De Lisle, J. (2005). "Reproductive ecology and evolution in the invasive terrestrial planarian Bipalium adventitium across North America". Pedobiologia. 49 (4): 367. doi:10.1016/j.pedobi.2005.04.002.
  7. Ogren, R. E.; Kawakatsu, M. (1987). "Index to the species of the genus Bipalium (Turbellaria, Tricladida, Terricola)". The Bulletin of Fuji Women's College Series 2. 25 (2): 79–119.
  8. Kawakatsu, M.; Ogren, R. E.; Froehlich, E. M.; Sasaki, G. (2002). "Additions and corrections to the previous land planarian indices of the world (Turbellaria, Seriata, Tricladida, Terricola) - 10". The Bulletin of Fuji Women's University. Ser. II. 40: 162–177.
  9. Stimpson, W. (1861). "On the genus Bipaliura". American Journal of Science and Arts. Series 2. 31: 134–135.
  10. Ogren, R. E. (1995). "Predation behaviour of land planarians". Hydrobiologia. 305 (1–3): 105–111. doi:10.1007/BF00036370.
  11. Ducey, P. K., M. McCormick, and E. Davidson. 2007. Natural history observations on Bipalium cf. vagum Jones and Sterrer, 2005 (Platyhelminthes: Tricladida), a terrestrial broadhead planarian new to North America. Southeastern Naturalist.
  12. Fiore, C.; Tull, J. L.; Zehner, S.; Ducey, P. K. (2004). "Tracking and predation on earthworms by the invasive terrestrial planarian Bipalium adventitium (Tricladida, Platyhelminthes)". Behavioural Processes. 67 (3): 327–334. doi:10.1016/j.beproc.2004.06.001. PMID 15518983.
  13. Ducey, P. K.; Messere, M.; Lapoint, K.; Noce, S. (1999). "Lumbricid Prey and Potential Herpetofaunal Predators of the Invading Terrestrial Flatworm Bipalium adventitium (Turbellaria: Tricladida: Terricola)". The American Midland Naturalist. 141 (2): 305. doi:10.1674/0003-0031(1999)141[0305:LPAPHP]2.0.CO;2.
  14. Dindal, D.L. (1970). "Feeding behavior of a terrestrial turbellarian Bipalium adventitium". The American Midland Naturalist. 83 (2): 635–637. doi:10.2307/2423970. JSTOR 2423970.
  15. Winsor, L. (1983). "A revision of the cosmopolitan land planarian Bipalium kewense Moseley, 1878 (Turbellaria: Tricladida: Terricola)". Zoological Journal of the Linnean Society. 79 (1): 61–100. doi:10.1111/j.1096-3642.1983.tb01161.x. ISSN 0024-4082.
  16. Ducey, P. K.; Cerqua, J.; West, L. J.; Warner, M. (2006). Eberle, Mark E (ed.). "Rare Egg Capsule Production in the Invasive Terrestrial Planarian Bipalium Kewense". The Southwestern Naturalist. 51 (2): 252. doi:10.1894/0038-4909(2006)51[252:RECPIT]2.0.CO;2.
  17. Stokes, A. N.; Ducey, P. K.; Neuman-Lee, L.; Hanifin, C. T.; French, S. S.; Pfrender, M. E.; Brodie, E. D.; Brodie Jr., E. D. (2014). "Confirmation and Distribution of Tetrodotoxin for the First Time in Terrestrial Invertebrates: Two Terrestrial Flatworm Species (Bipalium adventitium and Bipalium kewense)". PLoS ONE. 9 (6): e100718. Bibcode:2014PLoSO...9j0718S. doi:10.1371/journal.pone.0100718. PMC 4070999. PMID 24963791.
  18. Leal-Zanchet, A. M.; Baptista, V.; Campos, L. M.; Raffo, J. F. (2011). "Spatial and temporal patterns of land flatworm assemblages in Brazilian Araucaria forests". Invertebrate Biology. 130: 25–33. doi:10.1111/j.1744-7410.2010.00215.x.
  19. Jones, H. D. and B. Boag. 2001. The invasion of New Zealand flatworms. Glasgow Naturalist 23(supplement):77-83.
  20. Ogren, R.E. (1955). "Ecological observations on the occurrence of Rhynchodemus, a terrestrial turbellarian". Transactions of the American Microscopical Society. 74 (1): 54–60. doi:10.2307/3223842. JSTOR 3223842.
  21. Ducey, P. K.; Noce, S. (1998). "Successful invasion of New York State by the terrestrial flatworm, Bipalium adventitium". Northeastern Naturalist. 5 (3): 199–206. doi:10.2307/3858619. JSTOR 3858619.
  22. Esser, R. P. 1981. Land Planarians (Tricladida: Terricola). Contribution no. 227, Bureau of Nematology, Florida Department of Agricultural and Consumer Services, Division of Plant Industry, Gainesville, Fl.
  23. Hyman, L. H. 1951. The Invertebrates: Platyhelminthes and Rhynchocoela the acoelomate Bilateria, McGraw-Hill Book Co., London, 550 p.
  24. Choate, P.M., and Dunn, R.A., 1988. Land Planarians, Bipalium kewense Moseley and Dolichoplana striata Moseley (Tricladida: Terricola), IFAS Document EENY-049. Online:Available http://edis.ifas.ufl.edu/pdffiles/IN/IN20600.pdf
  25. Hyman, L.H. 1954. Some land planarians of the United States and Europe, with remarks on nomenclature. American Museum Novitates (1667): 1-21.
  26. Zaborski, E. 2002. Observations on feeding behavior by the terrestrial flatworm Bipalium adventitium (Platyhelminthes: Tricladida :Terricola) from Illinois. Am. Midl. Nat. 148: 401-408.
  27. Ogren, R.E. 1987. Description of a new three-lined planarian of the genus Bipalium (Turbellaria: Tricladida) from Pennsylvania, U.S.A. Trans. Am. Microsc. Soc. 106(1): 21-30.
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