Gigantorhynchus

Gigantorhynchus is a genus of Acanthocephala (thorny-headed worms, also known as spiny-headed worms) that parasitize marsupials, anteaters, and possibly baboons by attaching themselves to the intestines using their hook-covered proboscis. Their life cycle includes an egg stage found in host feces, a cystacanth (larval) stage in an intermediate host such as termites, and an adult stage where cystacanths mature in the intestines of the host. This genus is characterized by a cylindrical proboscis with a crown of robust hooks at the apex followed by numerous small hooks on the rest of the proboscis, a long body with pseudosegmentation, filiform lemnisci, and ellipsoid testes. The largest known specimen is the female G. ortizi with a length of around 240 millimetres (9.4 in) and a width of 2 millimetres (0.08 in). Genetic analysis on one species of Gigantorhynchus places it with the related genus Mediorhynchus in the family Gigantorhynchidae. Six species in this genus are distributed across Central and South America and possibly Zimbabwe. Infestation by a Gigantorhynchus species may cause partial obstructions of the intestines, severe lesions of the intestinal wall, and may lead to death.

Gigantorhynchus
Gigantorhynchus ortizi
Scientific classification
Kingdom: Animalia
Phylum: Acanthocephala
Class: Archiacanthocephala
Order: Gigantorhynchida
Family: Gigantorhynchidae
Genus: Gigantorhynchus
Hamann, 1892[1]

Taxonomy

The name Gigantorhynchus is derived from two Ancient Greek words: gígantas, which Otto Hamann used in 1892 as a descriptor for the family and genus when grouping the larger varieties of these worms,[lower-alpha 1] and rhúnkhos, meaning snout, nose, or beak, which describes the characteristic proboscis in this genus of Acanthocephala.[1] Phylogenetic analysis has been conducted on only one of the six species in the genus, G. echinodiscus, using the gene for 28S ribosomal RNA and confirms that this species forms a monophyletic group with the related genus Mediorhynchus in the family Gigantorhynchidae.[2] The type species is G. echinodiscus.[3]

Archiacanthocephala
Archiacanthocephala
Oligacanthorhynchidae
 

Macracanthorhynchus ingens

 
 

Oncicola venezuelensis

 
 

Oligacanthorhynchus tortuosa

 
 

Nephridiacanthus major

 
 
 
Moniliformidae

Moniliformis moniliformis

 
Gigantorhynchida
 

Mediorhynchus sp.

 
 

Gigantorhynchus echinodiscus

 
 
 
 
Phylogenetic reconstruction for select species in the class Archiacanthocephala[2][4]

Description

The genus Gigantorhynchus is characterized by the presence of a cylindrical proboscis with a crown of robust hooks at the apex followed by numerous small hooks on the rest of the proboscis.[2][5] The body, or trunk, is long with pseudosegmentation, the lemnisci are filiform, and the testes are ellipsoid.[2] Species of Gigantorhynchus are distinguished based on the number and size of hooks on the crown of the proboscis, the type of pseudosegmentation, and size of the ellipsoid eggs.[2] Males of all species possess eight cement glands which are used to temporarily close the posterior end of the female after copulation.[6] There is pronounced sexual dimorphism with the female often two or more times longer than the male.[2]

Species
G. echinodiscus
Measurements[2]Female (mm)Male (mm)
Length of proboscis0.49–0.710.45–0.65
Width of proboscis0.46–0.530.30–0.55
Length of proboscis receptacle0.63–0.740.48–0.64
Width of proboscis receptacle0.23–0.310.21–0.32
Length of trunk52.92–102.7914.80–45.29
Width of trunk0.79–1.130.53–0.99
Length of small rootless spines0.07
Length of lemnisci13.238.02 to 20.30
Size of anterior testis1.63–2.71 x 0.26–0.32
Size of posterior testis1.61–2.66 x 0.26–0.39
Size of eggs0.064 × 0.036
G. lopezneyrai
Measurements[2]Male (mm)
Length of proboscis1.131–1.5
Width of proboscis0.66
Length of trunk16–58
Width of trunk1–1.7
Length of lemnisci8
Size of anterior testis0.7–0.190
G. lutzi
Measurements[2][lower-alpha 2]Female (mm)Male (mm)
Length of proboscis1.695
Width of proboscis0.735
Length of trunk130–20035–60
Width of trunk1–2.50.75–1.15
Length of small rootless spines0.048
Length of lemnisci2.595
Size of anterior testis5.752–6.045 x 0.750–0.900
Size of eggs0.115 × 0.064
G. ortizi
Measurements[2][lower-alpha 3]Female (mm)Male (mm)
Length of proboscis1.45–1.72
Width of proboscis0.435–0.555
Length of proboscis receptacle0.750–0.920
Length of trunk130–24246–75
Width of trunk1.5–2.01.4–1.92
Length of small rootless spines0.05
Length of lemnisci5.48–6.80
Size of anterior testis1.98–3.0 x 0.56–0.96
Size of eggs0.079–0.085 × 0.049–0.054
G. pesteri
Measurements[2][lower-alpha 4]Female (mm)
Length of proboscis0.35
Width of proboscis0.1
Length of proboscis receptacle0.75
Width of proboscis receptacle0.18–0.2
Length of trunk15–18
Width of trunk0.8–0.9
Length of small rootless spines0.015
Length of lemnisci3.6–4
G. ungriai
Measurements[2][lower-alpha 5]Female (mm)Male (mm)
Length of proboscis0.189–1.0
Width of proboscis0.237–0.7
Length of trunk129–13622–36
Width of trunk1–1.60.78–1.58
Length of small rootless spines0.02–0.06
Length of lemnisci1.75–3.27
Size of anterior testis2.0–5.6 x 0.395–0.474
Size of eggs0.04–0.06 × 0.04

There are six valid species in the genus Gigantorhynchus,[3][7] although one species out of the six, G. pesteri, appears to be incorrectly assigned.[2]

G. echinodiscus is a tropical parasite of anteaters including the giant anteater (Myrmecophaga tridactyla), the southern tamandua (Tamandua tetradactyla) and the silky anteater (Cyclopes didactylus).[2] It has been found in Brazil,[2] Venezuela,[9] Panama,[10] and Trinidad Island.[11] Intermediate hosts include two species of termites from Brazil (Labiotermes emersoni and Orthognathotermes heberi).[12] It is the first species to be described in the genus Gigantorhynchus and is the type species.[3]

Morphological traits used to distinguish the species include a cylindrical proboscis with a crown having eighteen large hooks followed by 21 to 23 small rootless spines arranged in two longitudinal rows. The first row has six hooks measuring between 0.16 and 0.23 mm from the tip of the hook to the root. The second row has twelve hooks in pairs which are smaller than first row measuring between 0.18 and 0.19 mm from the tip of the hook to the root. The crown is separated from numerous small, rootless spines by a short space without hooks. Twenty-one to twenty-three small, rootless spines are arranged in longitudinal rows 0.05 to 0.08 mm long. The lemnisci reach the middle of the trunk and are sometimes bent back on themselves. Other traits include a lateral papilla at the base of the proboscis, a small region (2.24 to 3.21 mm long) after the proboscis with no segmentation, a ringed with no complete segmentation, large testes, eight cement glands in pairs occupying a region measuring between 0.98 and 2.13 mm long and between 0.45 and 0.76  mm wide, and a non-segmented region in the posterior end of the body.[2]

The male has two ellipsoid testes that are narrow and in tandem. The posterior end after the anterior testes without a segmented region measures between 5.45 and 8.53 mm. In the female, the gonopore is subterminal and the vagina has a sinuous lateral region shaped like a guitar. The genital pore including the vagina, uterus, and uterine bell is between 0.69 and 0.97 mm long. The eggs contain three membranes.[2]

  • Gigantorhynchus lopezneyrai Diaz-ungria, 1958[9]

G. lopezneyrai has been found parasitizing the Southern tamandua (Tamandua tetradactyla) in Venezuela.[2] The trunk is slightly segmented and no female measurements were taken in its original description. There are twelve hooks on the proboscis (4 in the first circle each around 0.235 mm long, and 8 in the second circle each around 0.106 mm long). The eight cement glands are organized in pairs.[2] Amato (2014) raised doubts about the validity of this species suggesting that the hook number and arrangement is an incorrect observation that needs to be revisited as no drawings of the proboscis showing the hook formation was published.[12] This species is named in honour of Carlos Rodríguez López-Neyra de Gorgot, a Spanish parasitologist.[9]

  • Gigantorhynchus lutzi Machado, 1941[13]

G. lutzi has been found parasitising a Bare-tailed woolly opossum (Caluromys philander) in Pará, Brazil[2] and infesting the small intestine of the Common opossum (Didelphis marsupialis) in Huanuco, Peru.[14] It was the second species in the genus Gigantorhynchus to be described. The body is ringed with no complete segmentation. There are twelve hooks on the proboscis including six in the first circle each measuring 0.285 by 0.165 mm, and six in the second circle measuring 0.225 by 0.135 mm. The eight cement glands are organized in pairs.[2]

  • Gigantorhynchus ortizi Sarmiento, 1954[15]

G. ortizi has been found infesting the intestines of the Brown four-eyed opossum (Metachirus nudicaudatus) and possibly a White-bellied slender opossum (Marmosops cf. noctivagus) both between Iquitos and Nauta in Peru.[16] Another survey found nearly 100% of the Brown four-eyed opossum were infected with this parasite in the Darien Province of Panama and the Departments of Chocó, Meta, and Nariño in Colombia. Infestations ranged from 20 to 60 worms per host almost completely obstructing the intestines in several of the sampled hosts.[17] The trunk pseudosegmented[15] or slightly segmented.[2] There are twelve hooks on the proboscis (six in the first circle each measuring 0.160 by 0.10 mm, and six in the second circle measuring 0.140 by 0.09 mm). The eight cement glands are organized in groups. It was named in honour of Dr. Javier Ortiz de La Puente, a Peruvian ornithologist from the Museum of the University of San Marcos, Lima, Peru who collected the brown four-eyed opossum from La Merced, Junin, Peru, which later was determined to have been infested with this new species of acanthocephalan.[15]

  • Gigantorhynchus pesteri Tadros, 1966[18]

G. pesteri was recorded from an unknown baboon species in Rhodesia (now Zimbabwe).[12][18] Gomes (2019) considers this Gigantorhynchus species to be incorrectly assigned due to a lack of information including missing registration number and deposit of specimen in a collection, missing type host species, unusual hook arrangement and number, and the description being based on only two immature females.[2] The proboscis has only four hooks whereas other species in the genus have either twelve or eighteen hooks on their proboscis.[18]

  • Gigantorhynchus ungriai Antonio, 1958[19]

G. ungriai has been found parasitizing a Southern tamandua (Tamandua tetradactyla) in Guayo, Delta Amacuro, Venezuela. The body is ringed and has a cylindrical shape with a complete segmentation consisting of a union in dorsal and ventral regions. The anterior end without segmentation measures 2 to 2.6 mm long. The retractable proboscis has eighteen hooks arranged in two circular rows. The first row has six hooks that are 0.140 to 0.2 mm long and the second row has twelve hooks that are 0.104 to 0.180 mm long. The eight cement glands occupy a space of 0.869 by 0.1896 mm.[2][19] The female genital tract made of an ovary-uterus extends throughout the length of the body. The male genitals occupy one quarter of the length of the body and contains elliptical testicles and eight peripheral prostate glands.[2][19] The species name was named after Carlos Díaz Ungría.[19]

Hosts

Gigantorhynchus species infest marsupials and myrmecophagids (anteaters) in Central and South America and possibly a baboon from Africa.[2] They are found in the intestines.[20] Cystacanths, the larval state of an Acanthocephalan, of G. echinodiscus were found in the hemocoels of soldier termites, the intermediate host. Termites infested with G. echinodiscus show head shape abnormalities and discolouration. Although worker termites feed directly on the feces of the final host (the location of the eggs of G. echinodiscus) it is not known why they are not infected with cystacanths or how they pass the eggs on to the soldiers where they develop into cystocanths. The giant anteater diet consists almost entirely of termites and the cystacanths from consumed termites mature and reproduce in the intestines of the host and the eggs are then passed in the feces. There are no paratenic hosts (hosts where parasites infest but do not undergo larval development or sexual reproduction) involved in the life cycle of G. echinodiscus.[12] Infestation of the giant anteater can cause at least partial obstructions of the intestines, severe lesions of the intestinal wall, and may lead to death.[21]

Notes
  1. "The larger varieties belong to this group/classification" (original German: "Wie der Name besagt, sind es große Formen, die hierher gehören.")[1]
  2. One value for the length of the proboscis and lemnisci, and width of proboscis in G. lutzi is reported for both sexes
  3. One range of values for the length of the proboscis and lemnisci, and width of proboscis in G. ortizi is reported for both sexes
  4. Data for G. pesteri reported from immature female specimens.
  5. One range of values for the length of the proboscis and lemnisci, and width of proboscis in G. ungriai is reported for both sexes
  6. A binomial authority in parentheses indicates that the species was originally described in a genus other than Gigantorhynchus. This species was originally named Echinorhynchus echinodiscus by Karl Moritz Diesing in 1851 but moved to Gigantorhynchus by Hamann in 1892.[3]

References
  1. Hamann, O. (1892). "Das System der Acanthocephalen" [The system of Acanthocephalans]. Zoologischer Anzeiger (in German). 15: 195–197. Retrieved 24 January 2020.
  2. Nascimento Gomes, Ana Paula; Cesário, Clarice Silva; Olifiers, Natalie; de Cassia Bianchi, Rita; Maldonado, Arnaldo; Vilela, Roberto do Val (December 2019). "New morphological and genetic data of Gigantorhynchus echinodiscus (Diesing, 1851) (Acanthocephala: Archiacanthocephala) in the giant anteater Myrmecophaga tridactyla Linnaeus, 1758 (Pilosa: Myrmecophagidae)". International Journal for Parasitology: Parasites and Wildlife. 10: 281–288. doi:10.1016/j.ijppaw.2019.09.008. PMC 6906829. PMID 31867208.
  3. Amin, O. M. (2013). "Classification of the Acanthocephala" (PDF). Folia Parasitologica. 60 (4): 275. doi:10.14411/fp.2013.031. PMID 24261131. Archived (PDF) from the original on 10 August 2017. Retrieved 27 September 2019.
  4. Amin, O.M.; Sharifdini, M.; Heckmann, R.A.; Zarean, M. (2020). "New perspectives on Nephridiacanthus major (Acanthocephala: Oligacanthorhynchidae) collected from hedgehogs in Iran". Journal of Helminthology. 94: e133. doi:10.1017/S0022149X20000073. PMID 32114988.
  5. Bhattacharya, S. B. (2007). Handbook on Indian Acanthocephala (PDF). Kolkata, Kinda: Director, Zool. Surv. India, Kolkata. pp. 14–15.
  6. Bush, Albert O.; Fernández, Jacqueline C.; Esch, Gerald W.; Seed, J. Richard (2001). Parasitism: the diversity and ecology of animal parasites. Cambridge, UK New York, NY: Cambridge University Press. p. 203. ISBN 0-521-66278-8. OCLC 44131774.
  7. "Gigantorhynchus Hamann, 1892". www.itis.gov. Integrated Taxonomic Information System (ITIS). 1 September 2019. Retrieved 1 September 2019.
  8. Diesing, Karl Moritz (1851). Systema helminthum vol. 2 (in Latin). v.2. Vindobonae:W. Braumüller. p. 512. doi:10.5962/bhl.title.46846.
  9. Diaz Ungría, C. D. (1958). "Sobre algunos Acantocefalos de Mammiferos venezolanos" [About some Acanthocephalans of Venezuelan mammals]. Reviews in Veterinary Medicine and Parasitology. 17: 191–204.
  10. Dunn, L. H. (1934). "Notes on the occurrence of Gigantorhynchus echinodiscus Diesing in the ant eater of Panama". Journal of Parasitology. 20 (4): 227–229. doi:10.2307/3272464. JSTOR 3272464.
  11. Cameron, Thomas W. M. (1939). "Studies on the endoparasitic fauna of Trinidad mammals: VI. Parasites of edentates". Canadian Journal of Research. 17d (12): 249–264. doi:10.1139/cjr39d-025.
  12. Amato, José F. R.; Cancello, Eliana M.; Rocha, Maurício M.; Carrijo, Tiago F. (2014). "Cystacanths of Gigantorhynchus echinodiscus (Acanthocephala, Gigantorhynchidae), in Neotropical Termites (Isoptera, Termitidae)". Neotropical Helminthology. 8 (2): 325–338. CiteSeerX 10.1.1.676.5918.
  13. Machado Filho, D. A. (1941). "Sobre alguns acantocéfalos provenientes do estado de Mato Grosso" [About some acanthocephalans from the state of Mato Grosso]. Revista Brasileira de Biologia (in Portuguese). 1 (1): 57–61.
  14. Tantaleán, Manuel; Sánchez, Lidia; Gómez, Luis; Huiza, Alina (2005). "Acantocéfalos del Perú" [Acanthocephalan from Peru]. Revista Peruana de Biología. 12 (1): 83–92. ISSN 1727-9933. Retrieved 9 June 2020.
  15. Sarmiento, Luz (August 1954). "Gigantorhynchus ortizi n. sp., an Acanthocephalan from Metachirus nudicaudatus". The Journal of Parasitology. 40 (4): 448–452. doi:10.2307/3273894. hdl:10919/40940. JSTOR 3273894. PMID 13192545. Retrieved 19 September 2020.
  16. Tantaleán, Manuel; Díaz, Mónica; Sánchez, Nofre; Portocarrero, Harold (2010). "Endoparásitos de micromamíferos del noroeste de Perú. 1: helmintos de marsupiales" [Endoparasites of micro-mammals from northwestern Peru. 1: marsupial helminths.]. Revista peruana de biología (in Spanish). 17 (2): 207–213.
  17. Thatcher, V. E.; Nickol, B. B. (1972). "Some acanthocephalans from Panama and Colombia" (PDF). Proceedings of the Helminthological Society of Washington. 39: 245–248. Retrieved 25 January 2020.
  18. Tadros, G. (1966). "On Gigantorhynchus pesteri n. sp from a Baboon". Journal of Helminthology. 40 (1–2): 181–186. doi:10.1017/S0022149X00034180. PMID 6006728.
  19. Antonio, H (1958). "Descripción de una nueva especie del género Gigantorhynchus Hamann, 1892 (Acanthocephala)" [Description of a new species of the genus Gigantorhynchus Hamann, 1892]. Acta Biologica Venezuelica (in Spanish). 2 (24): 291–298.
  20. Tantaleán, Manuel; Sánchez, Lidia; Gómez, Luis; Huiza, Alina (2005). "Acanthocephalan from Peru". Revista peruana de biología (in Spanish). 12 (1): 83–92. Retrieved 25 January 2020.
  21. Bomon, M. (2018). Management of gastrointestinal parasites in wildlife rehabilitation centers in Brazil (PDF) (Doctoral dissertation). Universidade de Lisboa, Faculdade de Medicina Veterinária. pp. 70–71. Retrieved 25 May 2020.

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