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Research Article A new genus and species of proteocephalidean tapeworm (Siluriformes: Auchenipteridae) from Brazil

Philippe Vieira Alves, Alain de Chambrier, Tomáš Scholz and José Luis Luque4

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Abstract: Frezella gen. n. is proposed to accommodate Frezella vaucheri1%5Tocantinsia piresi @+5>!5+&'!$'H +#+15!11^ 5!U1^5 ^'"51%%551 zone. Frezella can also be differentiated by having the internal longitudinal musculature hypertrophied laterally on both sides, the pres- !^15_1 canals situated slightly dorsomedian to ventral canals. Frezella vaucheriT. piresi, which &'H!+$' Keywords:^1515#5^5&'!+5>!5~

The Neotropical region hosts the highest diversity of Q RH+ piresi by @+3 @!5 5I!5HJ! !! (one of the principal southern tributaries of the Amazon "#- ! @ 5 @ 3M; 5 cephalidea). The highest diversity of tapeworm species was Tocantinsia Mees, to accommodate his new species T. de- recognised in large pimelodids such as Brachyplatystoma pressa @ H !5 ! Bleeker, Phractocephalus Agassiz, Pseudoplatystoma +&'!<!5 Bleeker and Zungaro $%&'# @ 3Q; 11 Tocantinsia ! + 1 depressa with Glanidium piresi, which actually does not many as 7–9 species of proteocephalideans (de Chambrier belong to Glanidium Lütken. Therefore, a new combina- 2333533356+ tion, T. piresi5+1@3Q;- 85;5&5;<!5- 1 ! R #1 ; teocephalidean cestodes also parasitise other groups of H+&'H! siluriforms, such as auchenipterids, in particular species + $ 5 + of Lacépède (see de Chambrier and Vaucher proteocephalidean cestodes are absent. 3335=#3 Morphological evaluation of the cestodes found in During studies on the parasitic helminths of freshwa- T. piresi >!!1- >!+1 resent a new species, which is described herein. In addition, &' ! $'5 the new species cannot be allocated to any of the known genera of proteocephalidean cestodes. Therefore, a new ge- Tocantinsia piresi @ +5 1 % nus is proposed to accommodate this new species.

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ZooBank number for article:""'+%"+"83&_88_;_$Q_;Q3;qRQ38q

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MATERIALS AND METHODS Type and only species: Frezella vaucheri sp. n. R +1 % ! +^ 11"H2 ++- UR'55+1- tory and then immediately dissected. Cestodes found were placed tematics and biology of proteocephalidean cestodes. The 5!^ name should be treated as femininum. hot 4% neutral formaldehyde solution and subsequently stored M‚6+;& Differential diagnosis. The new genus belongs to the the posterior end of the four remaining specimens was cut off # + 1 _ 38M33‚ „ 5 !1 + ^ Q~&15& 5! For morphology, specimens were stained with Mayer’s hydro- 3Q8533;H5+18- chloric carmine solution, dehydrated in an ethanol series, cleared era parasitising vertebrates of several groups (genera from !6+# ~ +1 %5 1 ++^5 Barsonella 6+5 '5 $ @^5 MX35ˆR^1- 3] ‹Brayela 5 3Q;] ‹Cangatiella #! ‚$+16- @ 5 33] ‹Euzetiella de Chambrier, + 25 333] Glanitaenia de Chambrier, Zeh- R @ +!5 52@^5;]‹Margaritaella Arredon- scoleces of the new species were prepared by the method de- = #5 ] ‹Proteocephalusˆ5 + +1  +! ‰ 3; QXQ]‹PseudocrepidobothriumU!!5] ~<1@5=! ‹Scholzia 6+5 = #5 X5 IŠI@8MRU ] Cairaella Coquille et de Chambrier, #15$66&55% $DJ!56'+@- Q] Crepidobothrium @5 3] Deblocktaenia ^16!13& 538]MacrobothriotaeniaR'538X]Ophio- are given in micrometres unless otherwise indicated. Abbrevia- taenia Š 5 3] Tejidotaenia R'5 38X5 "^M]M+ +]Thaumasioscolex &1"6

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Figs. 1–8. Frezella vaucheri gen. n. et sp. n. from Tocantinsia piresi$'Fig. 1. ^$> 8XM@<~=_#Š&HQ8MQ5!! Figs. 2, 3.^5!5!1]1$>8X1MU#6& 6_88]Figs. 4–7. 6^5 %5^'%5!1$>8XMU#6&6_88Fig. 8. Acicular !$>8XMU#6&6_88

%]! ^+1!- isolated pieces of strobila. Immature proglottids wider than %5 MXX R 5 5 "M5! 3]@M3%5XMMX "M8 5!1 ^MXMX;M5 + R ] much wider than neck (width of scoleces studied using R @MMQX]RM5^ #^+1^ & M3XM88 short, slightly folded, with subspherical muscular apical R553~%'! 5 XXM3 & 8XM3 5 58;XMXR53&^^5% %1R@- !5^' ^;% !

R#X58"8 # doi: 10.14411/fp.2015.006 Alves et al.: A new cestode genus (Proteocephalidea)

Figs. 9–11. Frezella vaucheri gen. n. et sp. n. from Tocantinsia piresi from Brazil. Fig. 9.^5!!]1$> 65f – CHIOC 37979a); note the presence of two distinct parts of the metascolex. Fig. 10.^5]1$>8X'M @<~=_#Š&HQ8M]+%Fig. 11. Pregravid proglottid, !!]1$>8X'M@<~=_#Š&HQ8MAbbreviations: ao – apical organ; cm – circular musculature; gc – glandular cells; lm – longitudinal musculature; mg – Mehlis’ gland; oc – osmoregulatory canals; od – oviduct. and density (Figs. 4–7). Proglottids covered with acicular _!R" RQ ;XM5XXMQX Inner longitudinal musculature well developed, formed M3M8H +1++ gravid proglottids. and ventral side, with conspicuous concentration of bun- Vas deferens strongly coiled, with loops forming elongat- dles on lateral sides of proglottids (Figs. 13–17). Ventral 5+5- osmoregulatory canals thin-walled, wide, median to lateral tid (Figs. 11, 14). Cirrus-sac elongate, thin-walled, slightly bands of vitelline follicles, slightly sinuous (Fig. 11). Dorsal widened towards distal (terminal) part (Fig. 12), 395–505 osmoregulatory canals thick-walled, relatively wide, sur- long and 100–160 wide (n = 31), its length representing rounded by vitelline follicles (Figs. 12–17). Another pair of MM‚^Q5 tiny, thin-walled canals situated slightly dorsomedian to ven- (internal vas deferens) strongly coiled (Fig. 12). Cirrus mus- tral canals, ventral to lateralmost lobes of ovary (Fig. 12); cular, occupies up to 60% (n = 30). Common genital atrium +!51 narrow, deep (Fig. 12). Genital pores alternating irregularly, Testes numerous, spherical, small, 50–70 in diameter, markedly pre-equatorial, situated at 23–41% (x = 31, n = 31) in 3 irregular layers (Figs. 13, 14), 279–420 in number of proglottid length from anterior margin (Fig. 11). ^XQ53H Ovary medullary, bilobed, follicular, with some dorsal by median line of proglottids (uterine stem) and vaginal ca- follicles penetrating between bundles of inner longitudinal

Folia Parasitologica 2015, 62: 006 Page 4 of 10 ";;{X8 &!"&#

Type locality: >!&5#5$- ' œ]X œˆ H1 "<1 68X' !+=$%+@QM33 M @<~=_#Š&H Q8M Abbreviations" M ] !M1+_%!1!]M] Remarks. Frezella vaucheri is the only species of the M_]M1]M monotypic Frezella and thus it is characterised by the fea- ]!M!1]!M! tures listed in its Differential diagnosis (see above). This ++1Tocantinsia piresi, a poorly %H- ^RX]!1 ture of the new species and the new genus is the pres- ;MM‚^585 ^%1 8XMMM‚^M5] 5„#U^ R@R+MM5- morphology, F. vaucheri somewhat resembles species of ;MM‚ !! Brooksiella 5 6++ #!5 3335 '6++X‚' %^- mature and pregravid proglottids. rounding the suckers (predominantly posterior to the suck- 21^5 5+^! 58Q‚;+- in Frezella356+;+ served) with asymmetrical chamber-like cavity near geni- In addition, Brooksiella praeputialis5- R5]!pars !53M;+1Mvs M copulatrix vaginae) surrounded by chromophilic cells, 5%1_!- ! R 2 J6+ M‚ 8‚ ;+ Peltidocotyle lenha ˆ5 3 ];_2 ^1 15515 ;6+2333- +1!!R5 + 6+ Š+Q;M3M‚^3‚ 'Q5++% Q;M3Q‚ ^  3‚ ++15#1ˆ- 5!1R 53; }15!1 Among the proteocephalideans that parasitise aucheni- 6+;5- H+5Nupelia tomasi de Chambrier gated longitudinal median concentration of chromophilic 25333+ galeatus cells in immature proglottids, with lumen appearing in last (Linnaeus) and Trachelyopterus cf. striatulus- immature proglottids. In mature proglottids, uterine stem #1+F. vaucheri in the possession of thick-walled, with median lumen and short diverticula. In ++- pregravid proglottids, uterine diverticula (lateral uterine culature concentrated on the lateral sides of the body (see branches) thin-walled, lined with few chromophilic cells. QQ;6+2333 } 1 1 +1 5 !^ slit-like pore. Q;6+2333<!5N. 5!8X]+- tomasi5 + +1 ~ #- 1+18MM53M !53333; 5+1%8MM subfamily), differs from the new species by numerous fea- Type host: Tocantinsia piresi @ + - 5+^5 "&5!Q R] %1+;;MQvsM3M; 1^&3 in F. vaucheri) and the smaller size of the body (total length ] ;MMvsM]^+X vs Q6+2333

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Fig. 13–18. Frezella vaucheri gen. n. et sp. n. from Tocantinsia piresi from Brazil. Fig. 13. Cross section at level of posterior part of ]1$>8X'M@<~=_#Š&HQ8MFigs. 14, 15.6!_!15!1]1 $>8XM68XM68XMM3M3Fig. 18. $>8XM@<~=_#Š&HQ8MAbbreviations"M]M_]M1]M- !]MJ1]M+1+1]M]M] M!]M!]M]!M!1]M++]M+ ]M]M]M]!M!]!M!]!M!]!M!] vo – ventral osmoregulatory canal.

Scholzia emarginata q5 QX +1 #- ture is much less developed in S. emarginata compared to ˆ5 3 5 the new species 6+XUE. ma- Phractocephalus hemioliopterus $5 lopteruri, + + & and Electrotaenia malopteruri (Fritsch, alongside the midline on the dorsal side of proglottids (see QQ8=@5335- 56+; 5Malapterurus electricus =5&! !^+1ˆ5 a peculiar arrangement of the inner longitutinal muscula- 3;5Endorchis piraeeba ˆ53; U5+ %%5 - laterally similarly as it is in F. vaucheri, but the muscula- mentosum Š5 #1-

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Table 1.#

+1 < Distribution Corallobothriinae Corallotaenia sp. @$ $%q3Q R'538X (Linnaeus) Endorchis auchenipteri osteomystax‹#$ 6+2333 ˆ53; 6+25333 @+ Endorchis sp. Trachelyopterus striatulus #$ 6+2333 Monticelliinae ‹ #$ 6+23335 Š53 6+25333 (Linnaeus), =#3 A. militaris Valenciennes Goezella danbrooksi Ageneiosus pardalis‹ @$ $%q3Q5 6+5@^5; (Linnaeus) 6+;+ Nupeliinae Nupelia tomasi Trachelyopterus galeatus‹ #$ 6+2333 #!533 6+25333 (Linnaeus), T. cf. striatulus #1 Luciaella ivanovae Ageneiosus inermis‹ #$ =#3 ˆ53; =#53 # Cangatiella arandasi Trachelyopterus galeatus‹ #$ #!@33 Š53 #!@533 Frezella vaucheri gen. n. et sp. n. A #1 Tocantinsia piresi* Zygobothriinae Gibsoniela mandube Ageneiosus inermis‹ &5#$ ˆ3X5 ˆ53 ˆ53X 3Q;5335 6+2333 Gibsoniela meursaulti Ageneiosus inermis‹, &5#$ ˆ3X5 6+25333 A. militaris 335 6+2333 ‹1]@$M@!+]#$M#!+]&M&' nae, e.g. Luciaella ivanovae = #5 3 monticelliaˆ53;MMRudolphiella the auchenipterid Ageneiosus inermis (Linnaeus), resem- szidati =#6+5Q3M8X ble the new cestode in the presence of some dorsal ovarian ˆ3;56+3356+ ^6+233M 23335=#6+ =#3<!5 The presence of a circular musculature in the suckers is species can be easily distinguished from F. vaucheri by the !^ !33; four types can be distinguished on the basis of its shape The presence of a chamber-like enlargement of the dis- 6+3U5 ^!5+! ^+1! in two thirds of proglottids in F. vaucheri, is rare among circular musculature only slightly developed, such as proteocephalidean cestodes. The terminal part of the va- Brooksiella praeputialis (Zygobothriinae) from the cetop- gina of Mariauxiella pimelodi 6+533X5 sid Cetopsis caecutiens^&'6 a parasite of Pimelodus ornatus  Pimelodus sp. Rudolphiella szidati =#6+5 $'55+11; ˆ5 3 Luciopimelodus 6+33X%F. vaucheri, pati2#~- which is often asymmetrical R5 3335=#6+ The precocious uterine aperture, which is present in 56+;+ F. vaucheri, has been reported in proteocephalideans para- sitising reptiles, such as species of Crepidobothrium Mon- DISCUSSION 5Q33TestudotaeniaR'538X5 ^ 1 1 %1 - eggs are laid immature through a longitudinal aperture in scription of genera or even subfamilies of proteocephali- !6+3QQ53Q35+5 5 & R'5 38] 6+3 6+ R'5 38X] = @5 335 The high number of the testes is a good discriminant MR'38X533;56+ ^1Thaumasioscolex didel- @1^^{ phidis6_='56+'5 absence of this structure was considered to be the most more testes than F. vaucheri ;MX;Qvs M3M; - #6_='& 333!1- other species from different subfamilies have a similar 6+;5<1 number of the testes, e.g. Proteocephalus paraguayensis X5'! 5 3M QM;; 5 Monticellia ventrei de character of this morphological structure, the presence of 6+25333MX5Nomimoscolex ad- ^5^-

R#X58"8 #M ";;{X8 &!"&# portance at least at the generic and species level (see, e.g. diagnosis. In addition, this diagnosis contains some errors, 6+233356„6- such as the presence of a spined cirrus (“Cirrus armed with +Q5' P]5- $%3Q;- 51^+Euzetiella tetraphylliformis 1 ^ #- 6+525333M6+ "+1' 333^+- (neck), e.g. Choanoscolex abscisus +5 Q38 J1 % X333]+1 acetabula indistinguishable from those in cyclophyllidean the tissue around each sucker, e.g. Brooksiella praeputia- cestodes). A putative synapomorphy, i.e. the presence of lis356+;+] simple gladiate microtriches, is also questionable and has to +1^+ +!+ of the suckers, e.g. Amphoteromorphus piraeeba ˆ- !+1+- 53;Q536+2 M& 33M ; H^Frezella vaucheri somewhat resem- ! +1%5 some of the ‘hooked’ tetraphyllidean-like cestodes such as 1^- those of Acanthobothrium !$5Q;3„- erated around the base of the suckers). In fact, this metas- 5^- ^+15 +J +1!^'5 1 + ! ^ with the anterior one strongly wrinkled and the posterior 11„^- 1 R & #- R+!5# teocephalidea, Spatulifer maringaensis #!5 is retained as a valid order in the present paper, but future 3Q3 and Peltidocotyle lenha have a somewhat similar systematic studies should focus on a comparative analysis ^5' ^1 As far as the authors are aware, there are no data on the 6; parasites of Tocantinsia piresi, which occurs in the Ama- 'H!+$5R Acknowledgements. The authors are indebted to Aldenice de ~' #5R+#R$# #1;H!+!- !+H% iour (feeds mainly on fruits and seeds) and its feeding be- also due to Jan Brabec for help with sequencing and analysing se- haviour is directly affected by the hydrologic cycle, i.e. wet „5+5% $DJ!& #' 16!3Q;5 =!@5I%#= 1\1- ~<1@5=! ration of its parasites. Frezella vaucheri thus represents the I <''}!R#M6- 1% &!H +- %I@^~- H+ <1@5=!H1+1 Based on a molecular phylogenetic study of tetraphyl- ‘Ciência sem fronteiras’ Brazilian program – visitant researcher _%+56; 1~X{1H$'}!- RI56~ proposed a new order, Onchoproteocephalidea, to accom- q!!6H6~#„I !^+!1 ŠŠ„~;M;MM{_5;X;{_Q5;88X{_ 11+$53 ]~R#$U~QQ838 #@53Q<!- QQQ5 U #1 51+ 28MM;;6'R#XX{{= morphological synapomorphies were provided in its brief #2&+16~#„

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&ǓǓdžDžǐǏDžǐ~I5Dždž6ljǂǎǃǓNJdžǓ&5=NJǍDždž#džǓǕNJdžǓǓǂ&& teocephalidea) from Pseudoplatystoma fasciatum#"- "&@- #!+&R# "#5Pseudoplatystoma fasciatum 8";8M;M #"#!$&5 &ǔlj &5 DŽljǐǍǛ H5 Dždž 6ljǂǎǃǓNJdžǓ &5 $ǓǂǃdžDŽ I5 Ǔǐǔ @5 with comments on microtriches of proteocephalideans. Folia ǂǓ#56ljǂǗǂǏ5@ǂǓNJǂǖǙI!Gan- #8";QMX8 gesia 6"#U1" &ǓǓdžDžǐǏDžǐ~I5=NJǍDždž#džǓǕNJdžǓǓǂ&&"Margaritaella 15#Š~ gracilis"#5 M";8; of Callichthys callichthys#"# $ǂǓǓǐǔ R$ "!^!- !+5&R#X3"33M8 '&MH@5&'5$ &ǓǓdžDžǐǏDžǐ~I5=NJǍDždž#džǓǕNJdžǓǓǂ&&5Dždž6ljǂǎǃǓNJdžǓ& &';"Q8M ;"&Pseudocrepidobothrium6"#-

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$Ǔǐǐnjǔq5qdžǂǓDžǐǓLJLJ5HŠ3Q"H- Dždž6ljǂǎǃǓNJdžǓ&5džLjǐ&&52ǂǖDŽljdžǓ6333"Euzetiella 6+5Nomimo- tetraphylliformis55"#5 scolex alovarius@"‰1+# parasite du poisson d’eau douce néotropical Paulicea luetkeni $ˆ;M"XM 5##8";M;M $Ǔǐǐnjǔq5ǂǔǎǖǔǔdžǏ=3Q;"# Dždž6ljǂǎǃǓNJdžǓ&5DŽljǐǍǛHQ"H6"#- 2' 5 @- Sorubimichthys planiceps #$ˆ3M"M;QMM8 "#&'!R#- 6ǂNJǓǂ I~5 IdžǏǔdžǏ 5 ˆǂdžǔDŽljdžǏǃǂDŽlj &5 ǍǔǐǏ #q5 ŠNJǕ- sitol. XX"MMQ ǕǍdžǘǐǐDžqHI;"M1- Dždž6ljǂǎǃǓNJdžǓ&5DŽljǐǍǛH5UǃǓǂljdždžǎ@<;"- !^1%1 scription of Electrotaenia malopteruriR5QQ86- UI#;;"XXMM " #5 Malapterurus electricus 6ǂDzdžDžǂ_=ǖǛǎǤǏ U65 Dždž 6ljǂǎǃǓNJdžǓ &5 DŽljǐǍǛ H " "@11#XM" Thaumasioscolex didelphidis"#- 3MM3 teocephalidae) from the black-eared opossum Didelphis marsu- Dždž6ljǂǎǃǓNJdžǓ&5DŽljǐǍǛH5ǖDŽljǕǂ;"H^ pialis@^5 ˆR6"#- I#.QM"83M8;M &'"+%- 6ǂǓǗǂǍljǐR@5DždždžǔdžǏDždž3Q;"&+ 1#QM"M3 Tocantinsia depressa5&&'- Dždž 6ljǂǎǃǓNJdžǓ &5 DŽljǐǍǛ H5 ǖDŽljǕǂ 5 #ǐǔdžǍ #5 Q"MMM @ǐǓǕdžǏǕljǂǍdžǓ@56ljǖǒǖNJǑNJǐǏDžǐ=ǖǂǓDžNJǂ68"H- Dždž6ljǂǎǃǓNJdžǓ&3QQ"Crepidobothrium garzonii n. sp. (Cestoda: 6" # &' # Bothrops alternatus Dum. Bibr. !#6#M"M ¼q5QX;"2#1! Dždž6ljǂǎǃǓNJdžǓ&52ǂǖDŽljdžǓ633M" ! Zool. 3X"8MM @ ˆ 3; ' Brachy- Dždž6ljǂǎǃǓNJdžǓ&3Q3"!Crepidoboth- ! Endorchis rium @5 3 6" # ˆ53;Nomimoscolexˆ53;1#- R ^UC. gerrardii$5Q8C. vi- M"3M peris$53!‰38"3MM Dždž6ljǂǎǃǓNJdžǓ&52ǂǖDŽljdžǓ6333"#@- Dždž6ljǂǎǃǓNJdžǓ&3Q3+"!Crepidoboth- "# rium @5 3 6" # R#1!R! R ^ Crepidobothrium dollfusi Freze, ^ !!‰8"8XM; 38X5Crepidobothrium lachesidis@653- Dždž6ljǂǎǃǓNJdžǓ&5‰džljǏDždžǓ@#52ǂǖDŽljdžǓ65@ǂǓNJǂǖǙI !‰ 38";XMQ ;"H!##15 Dždž6ljǂǎǃǓNJdžǓ&33" Proteocephalus para- + 115 guayensis53M6"# ! 1 # XM" Hydrodynastes gigasq5$+¼q5QX;#1 X3MM 1#8"QXM3M 6ljdžǓǗǚ Š 3" } 1 " Dždž6ljǂǎǃǓNJdžǓ&"&&'5Ama- a proposal from the international workshops on cestode system- zotaenia yvettae 5 5 " # MQR# X8"33M and 6ǐǒǖNJǍǍdž65Dždž6ljǂǎǃǓNJdžǓ&Q"Cairaella henrii gen. B. vaillanti#!‰ Q"M8 n. sp. n., a parasite of Norops trachyderma#15 Dždž6ljǂǎǃǓNJdžǓ&5&ǍǂǍǍǂnj~<5@ǂǓNJǂǖǙI"& Ophiotaenia nicoleae " #5 new tapeworm Postgangesia inarmata"#- a parasite of Thecadactylus rapicauda=%% " =5 Silurus glanis- R#XX"3MM8 U„= RǓǐdžǔdž5#ǂǖǍǚq) ;"R$ˆˆˆ+- 1#XX"33M3 +5+58{; Dždž6ljǂǎǃǓNJdžǓ&5$NJǏljHH5DŽljǐǍǛH"Ophiotaenia RǓdžǛdž2U38X"6122#- bungari n. sp. (Cestoda), a parasite of Bungarus fasciatus- R5&+~%5@5XQ "25 U]5U# relative ovarian size as a new and potentially useful diagnostic H538356~QX!\X3M 1#Q" =NJǍ Dždž #džǓǕNJdžǓǓǂ && 3" Luciaella ivanovae n. g., n. sp. 3MX " #" #15 Dždž 6ljǂǎǃǓNJdžǓ &5 6ǐǒǖNJǍǍdž 65 HnjǂDŽlj 25 @ǂǓNJǂǖǙ I Ageneiosus inermisŠ"&&- 3" Testudotaenia testudo @5 3; 1#M"MMQ "#5Apalone spinifera =NJǍDždž#džǓǕNJdžǓǓǂ&&5Dždž6ljǂǎǃǓNJdžǓ&"Rudolphiella Š"H1Amia calvaŠ#" szidati#"@5- Amiidae) in North America and erection of Testudotaeniinae n. linae) parasite of Luciopimelodus pati 25 Q; +1#M";3M8; #"#&+! Dždž6ljǂǎǃǓNJdžǓ&5džLjǐ&&33X"Mariauxiella pimelodi n. g., Rudolphiella lobosa+5Q3X!‰M" n. sp. (Cestoda: Monticelliidae): a parasite of pimelodid siluroid QM3X &1#"XMM8X <ǚǑȪǂ 25 ŠnjdžȦǯnjǐǗǤ &5 DŽljǐǍǛ H X"@1 Dždž6ljǂǎǃǓNJdžǓ&5džLjǐ&&5=NJǍDždž#džǓǕNJdžǓǓǂ&&X" and secondary structure characters in a reconstruction of phyl- "#- 15!_#- cephalidea), parasites of Phractocephalus hemioliopterus- #1"X3MM uriformes) from the Amazon and proposition of Scholzia gen. n. @dždžǔ =R 3M;" H & # - !‰"MXMMX #5~‰2"MX8 Dždž6ljǂǎǃǓNJdžǓ&5džLjǐ&&5@ǂǓNJǂǖǙI;+" @dždžǔ=R3Q;"&Tocantinsia#5~- of Brooksiella praeputialis and Goezeella siluri"#- 5&&'3"M; teocephalidea), parasites of Cetopsis coecutiens @NJǓǂǏDžǂNJǃdžNJǓǐ&3"#^^66 from the Amazon and proposition of Goezeella danbrooksi sp. ŠH @_=&- !‰ "M '5&^oX<~"‰XQ"MX

R#X58"8 #3 doi: 10.14411/fp.2015.006 Alves et al.: A new cestode genus (Proteocephalidea)

Ǔǐǔ@5DŽljǐǍǛH5<ǂǏǛdžǍǐǗǤ25@ǂDŽnjNJdžǘNJDŽnjI2010: džLjǐ&&56ljǖǃǃI65#ǂǗǂǏdžǍǍNJ=6333"6 ^1'611 &"%1+- #" scription of species. Rev. Bras. Zool. 16: 299–367. Folia Parasitol. 57: 37–46. džNJǔ"6!&- #ǂǗǂǏdžǍǍNJ=65@ǂDŽljǂDžǐDžǐǔǂǏǕǐǔ@<1991: Cangatiella U‰É+;M"8XMM arandasi56"#5 DŽljǎNJDžǕ=q3Q8"66<+%HU Parauchenipterus galeatus " & 66#U5$5R58MX Rio Parana. Rev. Bras. Biol. 7: 535–539. DŽljǐǍǛH5Dždž6ljǂǎǃǓNJdžǓ&5ǖDŽljǕǂ5ŠNJǕǕǍdžǘǐǐDžqHI5 džLjǐ&&3Q;"#&' ˆǂdžǔDŽljdžǏǃǂDŽlj&2013: 6" "1++1 #5+%Xenopeltis uni- Woodland as AnthobothriumH1&&';" color" ^ ! ! ‰^ 8;" ;QXM 86–94. 499. džLjǐ && 1992: Redescription of Gibsoniela mandube (Wood- DŽljǐǍǛH5Dždž6ljǂǎǃǓNJdžǓ&5@ǂǓNJǂǖǙI5ǖDŽljǕǂ2011: 53X6"#5Ageneio- Redescription of Corallobothrium solidum6"#- #"- cephalidea) and erection of a new genus, Essexiella, for tape- @U6' UI#3M";_ 87: 417–422. 1151. džLjǐ&&33;"#@53QU"ŠR- ˆǐǐDžǍǂǏDžˆ~R3;"&5 5&I&$1516# +15Myzophorus admon- 2+6&$U5ˆ5XMM3 ticellia, gen. and sp. n., parasitic in Pirinampus džLjǐ && 333" ^ 1 &'#18";M;3 ~@U ˆǐǐDžǍǂǏDž ˆ~R 3X" %+ 6'3;"MMX &'#1M"MMX

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Cite this article as: &!#256+&5'H5Š„IŠX"&- 65Tocantinsia piresi "&$'R Parasitol. 62: 006.

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