Dracunculus species

Cleveland, Christopher A., Garrett, Kayla B., Cozad, Rebecca A., Williams, Brianna M., Murray, Maureen H. & Yabsley, Michael J., 2018, The wild world of Guinea Worms: A review of the genus Dracunculus in wildlife, International Journal for Parasitology: Parasites and Wildlife 7 (3), pp. 289-300 : 290-293

2.1. Squamate Dracunculus species by region

2.1.1. Dracunculus ophidensis ( Brackett, 1938) and other species from North America

Dracunculus ophidensis was first described from garter snakes ( Thamnophis sirtalis ) from southern Michigan and subsequently found in

Species Host Geographic Anatomic location Prevalence (%) Species confirmation Reference location method

D. insignis River Otter ( Lontra Arkansas, USA Subcutaneous and intermuscular fascia of 12/184 (6.5) molecular Tumlison and Surf,

canadensis ) carpal and tarsal areas 2018

Racoon ( Procyon lotor ) Ontario, Canada NA NA molecular Elsasser et al., 2009

Ontario, Canada Subcutaneous tissues in the inguinal area, 253/553 morphology via male Crichton and thorax, abdomen and fascial layers of the (45.75) specimens Beverly-Burton, 1974 lower legs

Ontario, Canada Subcutaneous tissues in the inguinal area, left 1/1 (100) morphology via male Gibson and McKiel, and right axillary areas specimens 1972

Maryland, USA Subcutaneaous fascia of legs 1/1 (100) morphology via male Chitwood, 1950 specimens

D. lutrae River Otter Ontario, Canada NA NA molecular Elsasser et al., 2009

Ontario, Canada Subcutaneous tissues of thoracic, abdominal, 178/203 morphology via male Crichton and inguinal areas and intermuscular fascia of legs (87.7) specimens Beverly-Burton, 1974

Ontario, Canada Connective tissue beneath latissimus dorsi, NA morphology via male Crichton and subcutaneous tissues of thoracic, abdominal, specimens Beverly-Burton, 1973 inguinal areas and intermuscular fascia of legs

Dracunc- Badger ( Taxidea taxus ) Iowa, USA Subcutaneous and intermuscular fascia of 2/24 (8.3) NA Wittrock and Ulmer,

ulus carpal and tarsal areas 1974

sp. a Beaver ( Castor Kansas, USA Connective tissue beneath latissimus dorsi 2/63 (3) NA McKown et al., 1995

canadensis )

Fischer ( Martes pennanti ) New Hampshire, Subcutaneous and intermuscular fascia of 37/748 (4.9) NA Carlson and Vito,

USA carpal and metatarsal areas 1984

Marten ( Martes Ontario, Canada NA 1/405 (0.2) NA Seville and Addison,

americana ) 1995

Mink ( Neovison vison ) Ontario, Canada Connective tissue beneath latissimus dorsi, 14/42 (33) NA Schulte-Hostedde subcutaneous tissues of thoracic, abdominal, and Elsasser, 2011 inguinal areas and intermuscular fascia of legs

Arkansas, USA Subcutaneous and intermuscular fascia of 35/507 (6.9) NA Tumlison et al., 1984 carpal and metatarsal areas

Ohio, USA Subcutaneous and intermuscular fascia of 3/3 (100) NA Crites, 1963

carpal and metatarsal areas, tail musculature

Minnesota, USA Subcutaneous and intermuscular fascia of 3/3 (100) NA Huggins, 1958

carpal and metatarsal areas

New York, USA Intermuscular fascial layers of lower leg and NA NA Cheatum and Cook, ankles; subcutaneous position over pectoralis 1948

major

Minnesota, USA hind leg 2/72 (2.7) NA Erickson, 1946

Iowa, USA NA NA NA Benbrook, 1940

Wisconsin, USA NA NA NA Chaddock, 1940

Nebraska, USA NA NA NA Chitwood, 1933

Muskrat ( Ondatra Ontario, Canada Right inguinal region, left and right axillary 1/1 (100) NA Gibson and McKiel,

zibethicus ) regions 1972

Ontario, Canada NA NA NA Fyvie, 1969

Minnesota, USA Subcutaneous and intermuscular fascia of 1/1 (100) NA Huggins, 1958

carpal and metatarsal areas

Maryland, North NA NA NA Dikmans, 1948

Dakota, USA

Opossum ( Didelphis Maryland, USA Subcutaneous fascia 3/64 (4.7) NA Alexander et al.,

virginiana ) 1972

Otter New York, USA Intermuscular fascial layers of lower leg and NA NA Cheatum and Cook, ankles; subcutaneous position over pectoralis 1948

major

Racoon Florida, USA Subcutaneous and intermuscular fascia 9/54 (16.7) NA Keeling et al., 1993

Arkansas fascia of carpal and metatarsal areas 4/30 (13) NA Richardson et al .,

1992

Kentucky, USA Intermuscular fascial layers of lower leg and 10/70 (14.3) NA Cole and Shoop,

ankles 1987

Tennessee, NA 20/145 (13.8) NA Smith et al., 1985

Kentucky, USA

Illinois, USA Subcutaneous and intermuscular fascia of 2/245 (0.8) NA Snyder and carpal and metatarsal areas Fitzgerald, 1985

Arkansas Subcutaneous and intermuscular fascia of 35/507 (6.9) NA Tumlison et al., 1984 carpal and metatarsal areas

Georgia, USA NA 1/148 (0.7) NA Price and Harman, 1983

Connecticut, USA NA NA NA Diters and Ryan,

1980

Nebraska, Kansas, Subcutaneous and intermuscular fascia of 4/4 (100) NA Ewing and Hibbs ,

Missouri, USA carpal and metatarsal areas, facial areas 1966

North Carolina, Subcutaneous tissue and muscle fascia 14/209 (6.7), NA Harkema and Miller ,

South Carolina, 8/34 (23.5), 6/ 1964

Florida, USA 19 (31.6)

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Table 2 (continued)

Parasites identified as D. ophidensis have been reported from the mesentery of the Blackbelly garter snake ( Thamnophis melanogaster ) from several states in Mexico, indicating that this parasite may be widespread in garter snakes (Pérez-Ponce de León et al., 2001; Jiménez-Ruiz et al., 2002; Moravec, 2006). Dracunculus ophidensis also has been reported from northern water snakes ( Nerodia sipedon ) from Maryland, Minnesota, and Pennsylvania, and a plain-bellied water snake ( Nerodia erythrogaster ) from Michigan. However, confirmation in these other species requires molecular characterization or detection of male nematodes ( Mirza and Roberts, 1957; USNPC 1362086; Mirza, 1957; Moravec, 2006).

Unidentified Dracunculus spp . have been detected in a captive gopher snake ( Pituophis catenifer ) and a captive Florida kingsnake ( Lampropeltis getula floridana ) from The National Zoological Park (Washington D.C.) ( Mirza and Roberts, 1957; USNPC 1342730).

Minnesota ( Brackett, 1938). Nematodes were found in the subcutaneous tissue, serous membranes, and body cavity of infected snakes, and during summer months, gravid females were present in visible subcutaneous swellings ( Brackett, 1938; Moravec, 2006). Infections were only detected during the summer months, with signs of infection disappearing by fall or early winter ( Brackett, 1938). The arrangement of genital papillae and length of spicules in males are the most reliable characteristics used to differentiate D. ophidensis from other Dracunculus species. Spatial variation has been noted in D. ophidensis prevalence with a higher prevalence found in Minnesota compared with Michigan ( Brackett, 1938).

2.1.2. Dracunculus brasiliensis (Moravec and Santos, 2009) and other species from central and South America

A single subgravid female and part of a gravid female recovered from subcutaneous tissue and mesentery of one anaconda ( Eunectes murinus ) from Brazil were used to describe D. brasiliensis (Moravec and Santos, 2009) . Although most female Dracunculus spp . are morphologically similar, the shape of the female caudal end of D. brasiliensis differs from other Dracunculus spp . in that it is broad with a widelyrounded tip and the excretory pore located just posterior to the nerve ring. Additionally, the L1 of D. brasiliensis are considerably larger than most snake-infecting Dracunculus spp . (Moravec and Santos, 2009). Recently, D. brasiliensis was reported in a Brown-banded water snake ( Helicops angulatus ) from Matto Grosso state, Brazil (Quirino et al., 2018). Description of a male of this species is still needed.

Uncharacterized Dracunculus spp . have been reported from a number of free-ranging Central American, South American, and Caribbean snake species including free-ranging anaconda from Brazil and Venezuela, Pearl Island boa (Boa constrictor sabogae) from Panama, and boas (B. constrictor and reported as Corallus enydris ) from Trinidad (Muller, 1971; Calle et al., 1994; Moravec and Santos, 2009; USNM 1345047 and 1345052). In addition, unidentified Dracunculus nematodes have been recovered from several captive snakes that had been imported from South America including boas (B. constrictor and B. mexicana), an anaconda, and a bushmaster ( Lachesis muta ) ( Kotlan and Raitsits, 1923; Moravec, 1966; USNM 1338673 and 1335517).