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Revision of Corynosoma australe Johnston, 1937 (Acanthocephala: Polymorphidae) from a North American population using novel SEM images, Energy Dispersive X-ray Analysis, and molecular analysis


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Figs. 1, 2.

SEM of whole male (Fig. 1) and female (Fig. 2) specimens of Corynosoma australe from the intestines of Zalophus californianus in California. Note the continuity of the ventral field of spines from the neck to the genital area in the female which is covered with a copulation plug (Fig. 2).
SEM of whole male (Fig. 1) and female (Fig. 2) specimens of Corynosoma australe from the intestines of Zalophus californianus in California. Note the continuity of the ventral field of spines from the neck to the genital area in the female which is covered with a copulation plug (Fig. 2).

Figs. 3–8.

SEM of male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 3. Praesoma of a male worm showing the conical neck and the ventral spinose part of the anterior part of the trunk just anterior to the bulbus (white arrow) and the sensory pore (black arrow). 4. The anterior proboscis of a specimen showing the organization of the longitudinal rows of large hooks and the external appearance of the apical organ (arrow). 5. Apical view of the proboscis in Fig. 4 showing the external surface of the apical organ. 6. Lateral view of apical and subapical hooks. 7. Lateral view of middle hooks. 8. Posterior-most spine-like hooks. Note their arched curvature compared to more anterior hooks.
SEM of male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 3. Praesoma of a male worm showing the conical neck and the ventral spinose part of the anterior part of the trunk just anterior to the bulbus (white arrow) and the sensory pore (black arrow). 4. The anterior proboscis of a specimen showing the organization of the longitudinal rows of large hooks and the external appearance of the apical organ (arrow). 5. Apical view of the proboscis in Fig. 4 showing the external surface of the apical organ. 6. Lateral view of apical and subapical hooks. 7. Lateral view of middle hooks. 8. Posterior-most spine-like hooks. Note their arched curvature compared to more anterior hooks.

Figs. 9–14.

SEM of anterior bulbus and micropores of male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 9. Anterio-ventral view of the proboscis and bulbus of a female specimen. 10. Micropores at the anterior end of a proboscis. 11. Micropores on an anterior proboscis hook. 12. Micropores on anterior trunk spine. 13. Micropores at anterior end of a male trunk away from spines. 14. Micropores on the base of a posterior trunk spine in a male specimen.
SEM of anterior bulbus and micropores of male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 9. Anterio-ventral view of the proboscis and bulbus of a female specimen. 10. Micropores at the anterior end of a proboscis. 11. Micropores on an anterior proboscis hook. 12. Micropores on anterior trunk spine. 13. Micropores at anterior end of a male trunk away from spines. 14. Micropores on the base of a posterior trunk spine in a male specimen.

Figs. 15–20.

SEM of spines in male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 15. Anterior trunk spines in the bulbus area. 16. Transition area between bulbus spines (right) and posterior trunk spines. 17–19. Continuous ventral spines in one female specimen: 17. Midventral spines. 18. Posterior ventral spines. Note that the field of spines anterior to the genital spines constricts (arrow) but remains continuous with the genital spines without breaks. 19. Another, more detailed, perspective of posterior ventral spines of the same female in Fig. 17, 18 with arrow pointing to greatest constriction. Female gonopore is on the opposite, ventral, side (dorso-subterminal). 20. Posterior end of a male specimens with invaginated bursa revealing the organization of the genital spines there.
SEM of spines in male and female specimens of Corynosoma australe from the intestines of Zalophus californianus in California. 15. Anterior trunk spines in the bulbus area. 16. Transition area between bulbus spines (right) and posterior trunk spines. 17–19. Continuous ventral spines in one female specimen: 17. Midventral spines. 18. Posterior ventral spines. Note that the field of spines anterior to the genital spines constricts (arrow) but remains continuous with the genital spines without breaks. 19. Another, more detailed, perspective of posterior ventral spines of the same female in Fig. 17, 18 with arrow pointing to greatest constriction. Female gonopore is on the opposite, ventral, side (dorso-subterminal). 20. Posterior end of a male specimens with invaginated bursa revealing the organization of the genital spines there.

Figs. 21–26.

Detail of spines in specimens of Corynosoma australe from the intestines of Zalophus californianus in California using SEM and Gallium-cut sections. 21. SEM of an anterior trunk spine in the bulbus area. 22. A longitudinal Gallium-cut section of an anterior bulbus spine. Note the structure of the core element. 23. A longitudinal Gallium-cut section of a mid-trunk spine. 24. A longitudinal Gallium-cut section of a posterior spine. Note the structural differences between the outline and core of the spine types. 25. A dorso-terminal view of the posterior end of a male specimen showing the arrangement of the genital spines. Compare with Fig. 20. 26. A lateral view of a bursa showing its distal undulations and part of the genital spines (right).
Detail of spines in specimens of Corynosoma australe from the intestines of Zalophus californianus in California using SEM and Gallium-cut sections. 21. SEM of an anterior trunk spine in the bulbus area. 22. A longitudinal Gallium-cut section of an anterior bulbus spine. Note the structure of the core element. 23. A longitudinal Gallium-cut section of a mid-trunk spine. 24. A longitudinal Gallium-cut section of a posterior spine. Note the structural differences between the outline and core of the spine types. 25. A dorso-terminal view of the posterior end of a male specimen showing the arrangement of the genital spines. Compare with Fig. 20. 26. A lateral view of a bursa showing its distal undulations and part of the genital spines (right).

Figs. 27–32.

External orifices of male and female reproductive systems and eggs of Corynosoma australe from the intestines of Zalophus californianus in California using SEM and Gallium-cut sections. 27–29. The male bursa. 27. Bursa with the undulating lip, penis (white arrows), central cluster of prominent sensory bulbs (black arrows), and surrounding circles of specialized sensory domes (short black arrows). 28. The central part of a bursa showing the two types of sensory structures and the penis. 29. A higher magnification of the penis and the surrounding type of sensory domes. 30. The posterior end of a female showing the dorsal gonopore (right) and the continuity of the posterior ventral spines with the genital spines. 31. A smooth egg with no external ornamentation or dentations. 32. A cross Gallium-cut section of an egg showing 3 external egg shell membranes FM: fertilization membrane, IM: inner membrane, OM: outer membrane) with the fourth tightly enveloping the acanthor (A).
External orifices of male and female reproductive systems and eggs of Corynosoma australe from the intestines of Zalophus californianus in California using SEM and Gallium-cut sections. 27–29. The male bursa. 27. Bursa with the undulating lip, penis (white arrows), central cluster of prominent sensory bulbs (black arrows), and surrounding circles of specialized sensory domes (short black arrows). 28. The central part of a bursa showing the two types of sensory structures and the penis. 29. A higher magnification of the penis and the surrounding type of sensory domes. 30. The posterior end of a female showing the dorsal gonopore (right) and the continuity of the posterior ventral spines with the genital spines. 31. A smooth egg with no external ornamentation or dentations. 32. A cross Gallium-cut section of an egg showing 3 external egg shell membranes FM: fertilization membrane, IM: inner membrane, OM: outer membrane) with the fourth tightly enveloping the acanthor (A).

Figs. 33–36.

Line drawings of the female reproductive system and 3 selected hooks of Corynosoma australe from the intestines of Zalophus californianus in California. Fig. 33. The whole reproductive system of a 3.2 mm. long female. Note the presence of the Selective apparatus (SA) almost half way between the upper uterus (UU) connecting with the uterine bell (UB) and the lower uterus (LU) connecting with the dorsal vagina (V). Also note the dorsal and ventral short paravaginal ligament cords inserting in the body wall (PVBWLC) and the dorsal and ventral longer paravaginal ligament cords connecting with the uterine bell (PVUBLC). The para-vaginal muscular sheet wrapping around vagina can be observed. The continuity of trunk spines on the ventral side is evident. Fig. 34. A higher magnification of the posterior half of the reproductive system of a second female. Posterior muscular wrap not shown. Fig. 35. The 3 hook types: an anterior slender subapical hook (AH) with shorter root; a robust hook at the inflated part of the posterior proboscis (PH) with slightly longer root; and a small curved basal or near-basal hook (BH) with anteriorly directed root (manubrium). Fig. 36. The “uterine bell” of Corynosoma australe after Zdzitowiecki (1984, Fig. 2e, p. 362).
Line drawings of the female reproductive system and 3 selected hooks of Corynosoma australe from the intestines of Zalophus californianus in California. Fig. 33. The whole reproductive system of a 3.2 mm. long female. Note the presence of the Selective apparatus (SA) almost half way between the upper uterus (UU) connecting with the uterine bell (UB) and the lower uterus (LU) connecting with the dorsal vagina (V). Also note the dorsal and ventral short paravaginal ligament cords inserting in the body wall (PVBWLC) and the dorsal and ventral longer paravaginal ligament cords connecting with the uterine bell (PVUBLC). The para-vaginal muscular sheet wrapping around vagina can be observed. The continuity of trunk spines on the ventral side is evident. Fig. 34. A higher magnification of the posterior half of the reproductive system of a second female. Posterior muscular wrap not shown. Fig. 35. The 3 hook types: an anterior slender subapical hook (AH) with shorter root; a robust hook at the inflated part of the posterior proboscis (PH) with slightly longer root; and a small curved basal or near-basal hook (BH) with anteriorly directed root (manubrium). Fig. 36. The “uterine bell” of Corynosoma australe after Zdzitowiecki (1984, Fig. 2e, p. 362).

Figs. 37–39.

Energy Dispersive X-Ray spectra of Gallium cut anterior hook tip x-section (37), middle hook tip x-section (38), and posterior hook tip x-section (39) showing levels of sulfur in calcium and phosphorous in all hooks See Table 2 for more specific figures (bolded). Inserts: SEM of Gallium cut anterior hook tip cross-section (37), SEM of Gallium cut middle hook tip cross-section (38), and SEM of Gallium cut posterior hook tip cross-section (39). Note the high levels of calcium and phosphorous in anterior and middle hooks and the high level of sulfur in posterior hooks.
Energy Dispersive X-Ray spectra of Gallium cut anterior hook tip x-section (37), middle hook tip x-section (38), and posterior hook tip x-section (39) showing levels of sulfur in calcium and phosphorous in all hooks See Table 2 for more specific figures (bolded). Inserts: SEM of Gallium cut anterior hook tip cross-section (37), SEM of Gallium cut middle hook tip cross-section (38), and SEM of Gallium cut posterior hook tip cross-section (39). Note the high levels of calcium and phosphorous in anterior and middle hooks and the high level of sulfur in posterior hooks.

Fig. 40.

Phylogenetic relationships inferred using 18S gene sequences of Corynosoma australe and other acanthocephalan species. Nodal support from the two analyses is indicated as ML/BI and indicates values of bootstrap >70%. The scale-bar indicates the expected number of substitutions per site.
Phylogenetic relationships inferred using 18S gene sequences of Corynosoma australe and other acanthocephalan species. Nodal support from the two analyses is indicated as ML/BI and indicates values of bootstrap >70%. The scale-bar indicates the expected number of substitutions per site.

Fig. 41.

Phylogenetic reconstruction using mt Cox1 sequences of Corynosoma australe and sequences of other Acanthocephala deposited in the GenBank. The numbers indicate values of bootstrap >50%. Numbers above branches indicate nodal support as maximum likelihood (ML) and posterior probabilities from BI. The scale-bar indicates the expected number of substitutions per site.
Phylogenetic reconstruction using mt Cox1 sequences of Corynosoma australe and sequences of other Acanthocephala deposited in the GenBank. The numbers indicate values of bootstrap >50%. Numbers above branches indicate nodal support as maximum likelihood (ML) and posterior probabilities from BI. The scale-bar indicates the expected number of substitutions per site.

Fig. 42.

Median-joining haplotype networks based on Cox1 sequence data of Corynosoma australe. Each circle size represents the frequency of a haplotype as in the population.
Median-joining haplotype networks based on Cox1 sequence data of Corynosoma australe. Each circle size represents the frequency of a haplotype as in the population.

Fig. 43.

Phylogenetic tree from Maximum likelihood and Bayesian inference analysis of the Cox1 sequences of Corynosoma australe population. Outgroup: Corynosoma hannae. The scale-bar indicates the expected number of substitutions per site. Abbreviations used for isolates country: U1- present study isolates from USA, U- USA, M- Mexico, B- Brazil, A- Argentina, P- Peru. The color refers to isolates obtained as in the haplotype network (Fig. 42; red: USA, blue: Mexico, orange: Argentina, magenta: Brazil, aqua blue: Peru).
Phylogenetic tree from Maximum likelihood and Bayesian inference analysis of the Cox1 sequences of Corynosoma australe population. Outgroup: Corynosoma hannae. The scale-bar indicates the expected number of substitutions per site. Abbreviations used for isolates country: U1- present study isolates from USA, U- USA, M- Mexico, B- Brazil, A- Argentina, P- Peru. The color refers to isolates obtained as in the haplotype network (Fig. 42; red: USA, blue: Mexico, orange: Argentina, magenta: Brazil, aqua blue: Peru).

Chemical composition and localization of elements in hooks of Corynosoma australe from Zalophus californianus in California.

Anterior hooks Middle hooks Posterior hooks

Element* Tip x-section Middle Longitudinal section Tip x-section Tip edge Middle x-section Middle edge Tip x-section Middle edge
Magnesium (Mg) 1.21 0.23–1.67** 0.02–0.07 0.63 0.62 1.61 0.66 0.02 0.78–1.52
Sodium (Na) 0.25 0.00–0.04 0.00–0.03 0.00 0.03 0.08 0.00 0.02 0.05–0.07
Phosphorous (P) 9.95 15.74–18.71 12.78–20.35 10.64 11.00 21.32 16.10 2.82 14.91–20.49
Sulfur (S) 2.96 0.09–7.10 0.34–3.50 5.50 15.38 0.03 4.34 23.52 1.18–14.09
Calcium (Ca) 18.99 30.80–34.84 32.03–72.06 19.98 19.11 42.48 34.18 3.16 27.05–40.39

Chronological taxonomic history of Corynosoma australe from marine mammals, with special reference to ventral trunk spines in females.

Author Host Locality Described as Stage Ventral spines
Johnston (1937) Neophoca cinerea (Péron) South Australia C. australe Adults Discontinuous
Lincicome (1943) Zalophus californianis (Lesson) San Diego, California C. obtuscens Adults Continuous
Van Cleave (1953) Mycteroperca pardalis Gilbert Mazatalán, Mexico C. obtuscens Cystacanth Continuous
Morini & Boero (1960) Otaria flavescence Shaw Argentina C. otariae Adults Discontinuous
Zdzitowiecki (1984) Hydrurga leptonyx (Blainville) South Shetlands, Antarctica C. australe Adults 25% continuous
Smales (1986) Neophoca cinerea (Péron)Arctocephalus pusillatus Schreber South Australia C. australe Adults (Figs. 10,11) Discontinuous
Zdzitowiecki (1991) Few “suitable” definitive & paratenic hosts Antarctica C. australe Adults (Figs. 15 b, e) Discontinuous
Sardella et al. (2005) Arctocephalus australis (Zimmerman)Mirounga leonina (Linn.)Cynoscion guatucupa (Cuvier) Argentina C. australeC. australeC. australe AdultsAdultsCystacanths 85–100% continuous85–100% continuous85–89% continuous
Aznar et al. (2012) Otaria flavescens (Shaw) Argentina C. australe Adults (Fig. 1B) Discontinuous
Hernández-Orts et al. (2017a) Spheniscus magellanicus (Foster) Brazil C. australe Adults (Fig. 2B)Adults (Fig. 4B) Continuous & 82–89% continuous
Lisitsyna et al. (2018) Zalophus californianus (Lesson) SausalitoCalifornia C. obtuscens Adults Continuous
Lisitsyna et al. (2019) Zalophus californianus (Lesson) SausalitoCalifornia C. obtuscens & C. australe Adults 1% discontinuous
Present paper Zalophus californianus (Lesson) SausalitoCalifornia C. australe Adults Continuous with post. constriction

List of acanthocephalan species used for phylogenetic analysis based on the mt Cox1 gene sequences. Newly generated sequences are presented in bold.

Species Host Host origin GenBank accession nos. References
Corynosoma australe Zalophus californianus USA ON619618, ON614719 present study
Zalophus californianus USA MK119245–MK119249 Lisitsyna et al., 2019
Zalophus californianus Mexico MT676808–MT676818 García-Varela et al., 2020
Merluccius hubbsi Argentina MT676819–MT676822 García-Varela et al., 2020
Raneya brasiliensis Argentina MT676823–MT676824 García-Varela et al., 2020
Paralichthys isosceles Brazil KU314822 Fonseca et al., 2019
Stenella clymene Argentina MW724483 Hernández-Orts et al., 2021
Arctocephalus australis Argentina MF497333 Hernández-Orts et al., 2017a
Spheniscus magellanicus Brazil MF497335 Hernández-Orts et al., 2017a
Otaria flavescens Argentina KX957714, MF497334 Hernández-Orts et al., 2017a, b
Paralichthys adspersus Peru MZ920052–MZ920055 Mondragon-Martinez et al., 2021*
Paralabrax humeralis Peru MZ920056–MZ920059 Mondragon-Martinez et al., 2021*
Cheilodactylus variegatus Peru MZ920060–MZ920063 Mondragon-Martinez et al., 2021*
Otaria byronia Peru MZ920064–MZ920067 Mondragon-Martinez et al., 2021*
Corynosoma hannae Colistium guntheri New Zealand KX957724, KX957725 Hernández-Orts et al., 2017b
Leucocarbo chalconotus New Zealand KX957718–KX957721, KX957723 Hernández-Orts et al., 2017b
Phalacrocorax punctatus New Zealand KX957722 Hernández-Orts et al., 2017b
Phocarctos hookeri New Zealand KX957715–KX957717, JX442191 Hernández-Orts et al., 2017b; García-Varela et al., 2013
Peltorhamphus novaezeelandiae New Zealand KY909260–KY909263 Anglade & Randhawa, 2018
Corynosoma semerme Halichoerus grypus Germany MF001277 Waindok et al., 2018
Corynosoma semerme Callorhinus ursinus USA MK119253 Lisitsyna et al., 2019
Corynosoma obtuscens Halichoerus grypus New Zealand JX442192 García-Varela et al., 2013
Corynosoma villosum Callorhinus ursinus USA MK119251 Lisitsyna et al., 2019
Corynosoma validum Callorhinus ursinus USA MK119252 Lisitsyna et al., 2019
Corynosoma enhydri Enhydra lutris USA DQ089719 García-Varela & Nadler, 2006
Corynosoma magdaleni Phoca vitulina Germany MF078642 Waindok et al., 2018
Corynosoma nortmeri Phoca vitulina Germany MF001278 Waindok et al., 2018
Corynosoma strumosum Phoca vitulina USA EF467870 García-Varela & Pérez-Ponce de León, 2008
Pusa hispida botnica Finland EF467871 García-Varela & Pérez-Ponce de León, 2008
Zalophus californianus USA MK119250 Lisitsyna et al., 2019
Bolbosoma balaenae Balaenoptera physalus Italy MZ047281 Santoro et al., 2021
Bolbosoma turbinella Eschrichtius robustus USA JX442189 García-Varela et al., 2013
Andracantha phalacrocoracis Zalophus californianus USA MK119254 Lisitsyna et al., 2019
Hexaglandula corynosoma Nyctanassa violacea Mexico EU189488 Guillén-Hernández et al., 2008
Pseudocorynosoma anatarium Bucephala albeola Mexico KX688148 García-Varela et al., 2017
Pseudocorynosoma tepehuanesi Oxyura jamaicensis Mexico KX688139 García-Varela et al., 2017
Polymorphus obtusus Aythya affinis Mexico JX442195 García-Varela et al., 2013
Profilicollis bullocki Emerita analoga Mexico JX442197 García-Varela et al., 2013
Profilicollis chasmagnathi Oligosarcus jenynsii Argentina MT580124 Levy et al., 2020
Polymorphus trochus Fulica americana Mexico JX442196 García-Varela et al., 2013

Data for the population of Corynosoma australe used in the haplotype networking using mt Cox1 gene. Asterisk shows sequences unpublished on NCBI database. Newly generated sequences are presented in bold.

Geographical Locality* Cox1Haplotype no. ID in Fig. 46 GenBank accession nos. References
Pacific coast near San Francisco, California, USA H1 U1 ON619618, ON614719 Present study
Sausalito, California, USA H2 U MK119245-MK119249 Lisitsyna et al. 2019
Baja California, Mexico H2 M MT676816-MT676818 García-Varela et al. 2021
Rio de Janeiro, Brazil H3 B KU314822 Fonseca et al. 2019
Chubut, Argentina H4 A MW724483 Hernández-Orts et al. 2021
Baja California, Mexico H5 M MT676813 García-Varela et al. 2021
Sonora, Mexico H6 M MT676811 García-Varela et al. 2021
Baja California, Mexico H7 M MT676814 García-Varela et al. 2021
Sonora, Mexico H8 M MT676810 García-Varela et al. 2021
Northern Patagonia, Argentina H9 A MT676821 García-Varela et al. 2021
Sonora, Mexico H10 M MT676809 García-Varela et al. 2021
Baja California Sur, Mexico H11 M MT676808 García-Varela et al. 2021
Northern Patagonia, Argentina H12 A MT676823 García-Varela et al. 2021
Northern Patagonia, Argentina H13 A MT676819, MT676820, MT676822 García-Varela et al. 2021
Baja California, Mexico H14 M MT676815 García-Varela et al. 2021
Northern Patagonia, Argentina H15 A MT676824 García-Varela et al. 2021
Northern Patagonia, Argentina H16 A MF497333 Hernández-Orts et al. 2021
Baja California, Mexico H17 M MT676812 García-Varela et al. 2021
Rio de Janeiro, Brazil H18 B MF497335 Hernández-Orts et al. 2021
Northern Patagonia, Argentina H19 A MF497334 Hernández-Orts et al. 2021
Northern Patagonia, Argentina H20 A KX957714 Fonseca et al. 2019
Peru H21 P MZ920052, MZ920053, MZ920055, MZ920056-MZ920062, MZ920064-MZ920067 Mondragon-Martinez et al. 2021*
Peru H22 P MZ920054, MZ920063 Mondragon-Martinez et al. 2021*

Chemical composition of trunk spines and eggs of Corynosoma australe from Zalophus californianus in California.

Elements* Spines (longitudinal sections) Eggs (cross sections)


Anterior Middle Posterior Edge (shell) Center (acanthor)
Magnesium (Mg) 0.00 0.00 0.09 0.00 0.50
Sodium (Na) 0.00 0.00 0.00 0.00 0.00
Phosphorous (P) 1.39 0.00 1.16 0.91 7.73
Sulfur (S) 12.64 5.97 15.61 0 2.30
Calcium (Ca) 1.64 1.07 1.96 1.34 3.41

List of acanthocephalan species used for phylogenetic analysis based on the 18S rDNA gene sequences. Newly generated sequences are presented in bold, NA=host name not available.

Species Host Host origin GenBank accession nos. References
Corynosoma australe Zalophus californianus USA ON614192, ON614199 present study
Zalophus californianus USA MK119255 Lisitsyna et al., 2019
Phocarctos hookeri Mexico JX442168 García-Varela et al., 2013
Corynosoma obtuscens Callorhinus ursinus Mexico JX442169 García-Varela et al., 2013
Corynosoma validum Callorhinus ursinus Mexico JX442170 García-Varela et al., 2013
Corynosoma enhydri NA USA AF001837 Near et al., 1998
Corynosoma magdaleni Phoca hispida botnica Mexico EU267803 García-Varela et al., 2009
Corynosoma strumosum Phoca vitulina Mexico EU267804 García-Varela et al., 2009
Bolbosoma balaenae Nyctiphanes couchii Spain JQ040306 Gregory et al., 2011*
Bolbosoma turbinella Eschrichtius robustus Mexico JX442166 García-Varela et al., 2013
Bolbosoma vasculosum Lepturacanthus savala Indonesia JX014225 Verweyen et al., 2011
Andracantha gravida Phalacrocorax auritus Mexico EU267802 García-Varela et al., 2009
Hexaglandula corynosoma Nyctanassa violacea Mexico EU267808 García-Varela et al., 2009
Pseudocorynosoma anatarium Bucephala albeola Mexico EU267801 García-Varela et al., 2009
Pseudocorynosoma constrictum Anas clypeata Mexico EU267800 García-Varela et al., 2009
Profilicollis bullocki Emerita analoga Mexico JX442174 García-Varela et al., 2013
Profilicollis botulus Somateria mollissima Mexico EU267805 García-Varela et al., 2009
Polymorphus trochus Fulica americana Mexico JX442196 García-Varela et al., 2013
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