http://folia.paru.cas.cz This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Phylogenetic relationships and systematics of tapeworms of the family Davaineidae (Cestoda, Cyclophyllidea), with emphasis on species in rodents Voitto Haukisalmi1 , Alexis Ribas2, 3 , Jean-Pierre Hugot4 , Serge Morand5 , Kittipong Chaisiri6, Kerstin Junker7 , Sonja Matthee8 , Andrea Spickett7, 8 , Jukka T. Lehtonen9, Carlos Feliu10 and Heikki Henttonen11 1 Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, Helsinki University, Finland; 2 Department of Biology, Health and Environment Faculty of Pharmacy and Food Science, University of Barcelona, Spain; 3 Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Spain; 4 Museum National d’Histoire Naturelle, Département Origine et Evolution, Institut de Systématique, Evolution et Biodiversité, Paris, France; 5 IRL HealthDEEP, CNRS - Kasetsart University - Mahidol University, Bangkok, Thailand; 6 Department of Helminthology, Faculty of Tropical Medicine, Mahidol University, Bangkok, Thailand; 7 National Collection of Animal Helminths, Epidemiology, Parasites and Vectors Programme, ARC-Onderstepoort Veterinary Institute, Onderstepoort, South Africa; 8 Department of Conservation Ecology and Entomology, Stellenbosch University, Matieland, South Africa; 9 Faculty of Biological and Environmental Sciences, Helsinki University, Finland; 10 Faculty of Pharmacy, University of Barcelona, Spain; 11 Natural Resources Institute Finland (Luke), Helsinki, Finland Abstract: The present study aims at clarifying the poorly known phylogenetic relationships and systematics of cestodes of the family Davaineidae Braun, 1900 (Cyclophyllidea), primarily the genus Raillietina Fuhrmann, 1920 and of the subfamily Inermicapsiferinae (Anoplocephalidae) from mammals (mostly rodents, 31 new isolates) and birds (eight new isolates). Phylogenetic analyses are based on sequences of the large subunit ribosomal RNA gene (28S) and mitochondrial NADH dehydrogenase subunit 1 gene (nad1). The main phylogenetic pattern emerging from the present analysis is the presence of three independent lineages within the main clade of the subfamily Davaineinae, one of which is almost entirely confined to species from rodents and the other two show a mixture of species from birds and mammals. It is suggested that the major diversification of the main clade took place in birds, possibly in galliforms. The subsequent diversification included repeated host shifts from birds to mammals and to other birds, and from rodents to other mammals, showing that colonisation of new host lineages has been the main driver in the diversification of davaineine cestodes. It is also shown that all isolates of Inermicapsifer Janicki, 1910, mainly from rodents, form a monophyletic group positioned among Raillietina spp. in the “rodent lineage”, indicating that the genus Inermicapsifer is a member of the family Davaineidae. This means that the subfamily Inermicapsiferinae and the family Inermicapsiferidae should be treated as synonyms of the Davaineidae, specifically the subfamily Davaineinae. Three additional genera generally included in the Inermicapsiferinae, i.e. Metacapsifer Spasskii, 1951, Pericapsifer Spasskii, 1951 and Thysanotaenia Beddard, 1911, are also assigned here to the Davaineidae (subfamily Davaineinae). Raillietina spp. were present in all three main lineages and appeared as multiple independent sublineages from bird and mammalian hosts, verifying the non-monophyly of the genus Raillietina and suggesting a presence of multiple new species and genera. Keywords: Davaineinae, Anoplocephalidae, Raillietina, Inermicapsifer, Inermicapsiferinae, phylogeny, 28S, nad1 Research Article Address for correspondence: V. Haukisalmi, Koukkurannankatu 3 B 37, FIN-33870 Tampere, Finland. Phone: +358 50 4645058; E-mail: haukisal@ protonmail.com Institute of Parasitology, Biology Centre CAS Folia Parasitologica 2024, 71: 011 doi: 10.14411/fp.2024.011 Cestodes of the family Davaineidae Braun, 1900 (Cy- clophyllidea) comprise a species-rich group with a world- wide distribution in birds and mammals. The family includes at least 450 nominal species and 37 genera (Mari- aux et al. 2017). Approximately 80 nominal species have been described from mammals, of which ca. 35 species doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 2 of 11 are from rodents (Movsesyan 2003a,b, Caira et al. 2024). The Davaineidae is currently considered to include two subfamilies, i.e. Davaineinae and Idiogeninae, although Ophryocotylinae and Cotugniinae have sometimes been regarded as valid subfamilies. For the most recent over- view of morphologic features and classification of davain- eids, see Mariaux et al. (2017). The monophyly of the family, to the exclusion of the genus Ophryocotyle Friis, 1870, was supported by the mo- lecular phylogenetic analysis of Waeschenbach and Little- wood (2017), part of the Planetary Biodiversity Inventory project “Tapeworms from vertebrate bowels of the Earth” (Caira and Jensen 2017). The monophyly of the Davain- eidae, including Ophryocotylinae, was also supported by the phylogenetic analysis of Hoberg et al. (1999) based on morphologic characters. The phylogenetic position of the Davaineidae with respect to other cyclophyllidean families remains undefined, although it is probably a relatively ear- ly diverging group (Waeschenbach and Littlewood 2017). However, the analysis of Hoberg et al. (1999) suggested a derived phylogenetic position for the Davaineidae. The phylogenetic relationships between species and genera of davaineid cestodes are largely unknown. The existing studies have dealt with very limited assemblages, including mainly species of Raillietina Fuhrmann, 1920 from birds (O’Callaghan 2004, Littlewood et al. 2008, Butboonchoo et al. 2016, Siddiqui et al. 2023). Individual DNA sequences of Raillietina species from mammals (ro- dents, pangolin) have been published by Oliveira Simões et al. (2017), Tuli et al. (2022) and Panti-May et al. (2023). In the latter studies, the species from mammals were po- sitioned among species from birds, suggesting that they originated as a consequence of a shift from the latter hosts. The genus Raillietina, of the subfamily Davaineinae, is by far the most species-rich genus of davaineid cestodes, with ca. 200 nominal species, of which ca. 165 species in birds and ca. 35 species in mammals, most of the latter spe- cies (22) in rodents (Movsesyan 2003a, Caira et al. 2024). The prevailing taxonomic concept of the genus Raillietina is, however, very wide and evidently artificial, including all species with unilateral genital pores and multiple eggs in each parenchymatous capsule (Jones and Bray 1994, Movsesyan 2003a). Not surprisingly, the molecular phy- logenetic analysis of Littlewood et al. (2008), based on six species of Raillietina and Fuhrmannetta malakartis Ma- hon, 1958 from birds, strongly suggested that the former genus is not monophyletic. However, large-scale phyloge- netic analyses that would allow us to consider new classi- fication schemes for the genus Raillietina are still lacking. The cestodes of the Inermicapsiferinae, a subfami- ly traditionally assigned to the family Anoplocephalidae (see Spasskii 1951, Schmidt 1986 and Beveridge 1994), are morphologically very similar to the davaineid ces- todes, particularly Raillietina, the absence of a rostellum and armature in the former group being the only clear-cut morphologic difference between these taxa. Lopez-Neyra (1954, 1955) transferred several davaineid genera to the Anoplocephalidae, while Baer and Fain (1955) transferred inermicapsiferines (then included in the Linstowiidae) to the subfamily Davaineinae. The latter action was support- ed by Mettrick and Weir (1963) and also later by Spasskii (1996). However, the monophyly and systematic position of inermicapsiferine cestodes has not been tested using molecular phylogenetic methods. The present study aims at clarifying the poorly known phylogenetic relationships of davaineid cestodes of birds and mammals (particularly rodents) and inermicapsiferine cestodes of rodents using phylogenetic analyses based on nuclear and mitochondrial DNA sequences. The observed phylogenetic relationships are used to infer patterns of di- versification and systematic relationships in the Davainein- ae, particularly in the genera Raillietina and Inermicapsifer Janicki, 1901. MATERIALS AND METHODS Material The present material includes 39 new isolates of davain- eid and inermicapsiferine cestodes from mammals (31 isolates, mostly from rodents) and birds (eight isolates) (Table 1). The iso- lates were assigned to the genera Raillietina, including the type species Raillietina tetragona (Molin, 1858) from the domestic fowl, Paroniella Fuhrmann, 1920, Delamuretta Spasskii, 1977, Calostaurus Sandars, 1957, Skrjabinia Fuhrmann, 1920, Cotug- nia Diamare, 1893 and Inermicapsifer according to the keys of Jones and Bray (1994) and Beveridge (1994). Species-level iden- tification was based on the descriptions available in Movsesyan (2003a,b) and in Caira et al. (2024). However, most of the isolates of Raillietina and Inermicapsifer from rodents could not be iden- tified to species. The present material includes three isolates from Cape Verde, which were treated as Thysanotaenia congolensis Dronen, Simcik, Scharninghausen et Pitts, 1999 (Inermicapsiferi- nae) by Świderski et al. (2015a,b) and Miquel et al. (2016). How- ever, they are placed here in the genus Inermicapsifer (species “4”) due to a lack of significant morphologic differences between the present specimens from Cape Verde and Africa, and the Iner- micapsifer species described from African rodents (Caira et al. 2024). For the host species and other background information of the present isolates, see Table 1. Molecular and phylogenetic analysis The present molecular analysis utilises sequences of the large subunit ribosomal RNA gene (28S) and mitochondrial NADH dehydrogenase subunit 1 gene (nad1). For 28S, DNA was am- plified using two alternative pairs of primers: (i) XZ-1 (forward, 5’-ACCCGCTGAATTTAAGCATAT-3’) of Waeschenbach et al. (2007), which differs from the original XZ-1 of Van der Auwera et al. (1994) by having one ‘‘Y’’ replaced with ‘‘T’’ (in bold), and 1500R (reverse, 5’-GCTATCCTGAGGGAAACTTCG-3’) of Littlewood et al. (2008) (ca. 1500 bp), and (ii) U178 (for- ward, 5’-GCACCCGCTGAAYTTAAG-3’) and L1642 (reverse, 5’-CCAGCGCCATCCATTTTCA-3’ (ca. 1500 bp), both from Lockyer et al. (2003). For nad1, DNA was amplified with primers Cyclo_Nad1F (forward, 5’-GGNTATTSTCARTNTCGTAAG- GG-3’) and Cyclo_trnNR (reverse, 5’-TT CYTGAAGTTAA- CAGCATCA-3’) (ca. 800 bp) of Littlewood et al. (2008). Standard 50 μl PCR was performed using cycling conditions of Waeschenbach et al. (2007) and Littlewood et al. (2008) for doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 3 of 11 Ta bl e 1. G en Ba nk a cc es sio n nu m be rs , v ou ch er n um be rs a nd b ac kg ro un d in fo rm at io n fo r i so la te s us ed in th e pr es en t p hy lo ge ne tic a na ly se s. Th e po sit io n of C al os ta ur us s pp . a nd R ai lli et in a sp p. (M ex ic o) w it hi n th e cl ad e B is r el at iv el y po or ly s up po rt ed ( F ig . 1 ). D ep os it or ie s of v ou ch er s: “ M Z B ”, M us eu d e Ci èn ci es N at ur al s d e Ba rc el on a, S pa in ; ” FM N H ”, F in ni sh M us eu m o f N at ur al H is- to ry ; “ C N H E ”, C ol ec ci ón N ac io na l d e H el m in to s, M ex ic o; “ S A M ”, S ou th A us tr al ia n M us eu m . Ce sto de sp ec ie s D N A co de H os t s pe ci es H os t f am ily Co un try Lo ca lit y Vo uc he rs G en Ba nk (2 8S ) G en Ba nk (n ad 1) So ur ce Cl ad e A Ra ill ie tin a sp . 1 D V 7 Ba nd ic ot a in di ca M ur id ae Vi et na m H on g G nu M Z B 2 02 4- 25 84 - O R7 95 87 8 Pr es en t s tu dy Ra ill ie tin a sp . 2 D N 9 Ra ttu s e xu la ns M ur id ae Vi et na m Ca o La nh M Z B 2 02 4- 25 83 - O R7 95 87 7 Pr es en t s tu dy Ra ill ie tin a sp . 3 BU 4 Le op ol da m ys sa ba nu s M ur id ae Th ai la nd Lo ei M Z B 2 02 4- 25 77 - O R7 95 87 0 Pr es en t s tu dy Ra ill ie tin a sp . 3 CI 3 Ra ttu s t an ez um i M ur id ae La os Lu an g Pr ab an g M Z B 2 02 4- 25 78 - O R7 95 87 5 Pr es en t s tu dy Ra ill ie tin a sp . 3 C Z 0 B. in di ca M ur id ae La os La k M Z B 2 02 4- 25 79 - O R7 95 87 6 Pr es en t s tu dy Ra ill ie tin a sp . 4 D A 4 Le op ol da m ys . h er be rt i M ur id ae La os La k M Z B 2 02 4- 25 80 O R8 05 58 6 K P1 71 52 7 Pr es en t s tu dy Ra ill ie tin a sp . 5 W 54 N es om ys ru fu s N es om yi da e M ad ag as ca r Ra no m af an a FM N H K N .4 34 40 K P1 71 52 3 K P1 71 52 6 Pr es en t s tu dy Ra ill ie tin a sp . 6 BV 3 N es om ys a ud eb er ti N es om yi da e M ad ag as ca r Ra no m af an a FM N H K N .4 34 41 O R8 05 58 1 - Pr es en t s tu dy Ra ill ie tin a sp . 7 BT 4 La on as te s a en ig m am us D ia to m yi da e La os K ha m m ou an e FM N H K N .4 34 37 O R8 05 57 9 O R7 95 86 7 Pr es en t s tu dy Ra ill ie tin a sp . 8 BT 5 L. a en ig m am us D ia to m yi da e La os K ha m m ou an e FM N H K N .4 34 39 O R8 05 58 0 O R7 95 86 8 Pr es en t s tu dy Ra ill ie tin a sp . 9 BT 2 L. a en ig m am us D ia to m yi da e La os H in bo um FM N H K N .4 34 38 O R8 05 58 3 O R7 95 86 9 Pr es en t s tu dy Ra ill ie tin a sp . C Z 8 L. h er be rt i M ur id ae La os La k M Z B 2 02 4- 25 82 O R8 05 58 5 - Pr es en t s tu dy D el am ur et ta sp . BU 5 L. sa ba nu s M ur id ae Th ai la nd Lo ei - - O R7 95 87 1 Pr es en t s tu dy D el am ur et ta sp . CI 1 R. ta ne zu m i M ur id ae La os Lu an g Pr ab an g FM N H K N .4 34 44 , M Z B 2 02 4- 25 81 O R8 05 58 4 O R7 95 87 4 Pr es en t s tu dy In er m ic ap si fe r s p. 1 EP 3 Le m ni sc om ys ro sa lia M ur id ae So ut h A fri ca M oo in oo i - O R8 05 58 9 O R7 95 88 6 Pr es en t s tu dy In er m ic ap si fe r s p. 1 ES 4 L. ro sa lia M ur id ae So ut h A fri ca Z ee ru st FM N H , K N .4 34 49 , K N .3 64 8 O R8 05 59 0 O R7 95 88 2 Pr es en t s tu dy In er m ic ap si fe r s p. 2 ES 9 Ae th om ys c hr ys op hi lu s M ur id ae So ut h A fri ca M oo in oo i - - O R7 95 88 4 Pr es en t s tu dy In er m ic ap si fe r s p. 2 ET 5 A. c hr ys op hi lu s M ur id ae So ut h A fri ca Z ee ru st FM N H K N .4 34 48 - O R7 95 88 5 Pr es en t s tu dy In er m ic ap si fe r s p. 3 EI 4 Sm ut si a te m m in ck ii M an id ae So ut h A fri ca - - - O R7 95 88 1 Pr es en t s tu dy In er m ic ap si fe r s p. 4 CY 0 Ra ttu s r at tu s M ur id ae Ca pe V er de Sa nt ia go Is la nd - - K P1 71 52 8 Pr es en t s tu dy In er m ic ap si fe r s p. 4 CY 7 R. ra ttu s M ur id ae Ca pe V er de Sa nt ia go Is la nd FM N H K N .4 34 47 K P1 71 52 2 - Pr es en t s tu dy In er m ic ap si fe r s p. 4 CY 2 M us m us cu lu s M ur id ae Ca pe V er de Sa nt ia go Is la nd FM N H K N .4 34 46 K P1 71 52 1 K P1 71 52 9 Pr es en t s tu dy In er m ic ap si fe r s p. 4 ES 5 M as to m ys c ou ch a M ur id ae So ut h A fri ca K aa lp la as FM N H K N .3 64 7 O R8 05 59 1 O R7 95 88 3 Pr es en t s tu dy In er m ic ap si fe r s p. 5 FB 3 Rh ab do m ys d ile ct us M ur id ae So ut h A fri ca A lic e, E as t C ap e FM N H K N .3 64 9 O R8 05 59 4 - Pr es en t s tu dy Cl ad e B Ra ill ie tin a co re en si s H on da , 1 93 9 CA 7 Ap od em us a gr ar iu s M ur id ae K or ea D M Z 1 FM N H K N .4 34 34 K P1 71 52 5 K P1 71 53 1 Pr es en t s tu dy R. e ch in ob ot hr id a (M eg ni n, 1 88 0) EV 0 G al lu s g al lu s Ph as ia ni da e Th ai la nd U do n Th an i - - O R7 95 88 7 Pr es en t s tu dy R. te tr ag on a (M ol in , 1 85 8) EV 1 G . g al lu s Ph as ia ni da e Th ai la nd U do n Th an i - O R8 05 59 2 O R7 95 88 8 Pr es en t s tu dy R. te tr ag on a EV 6 G . g al lu s Ph as ia ni da e Th ai la nd U do n Th an i - O R8 05 59 3 O R7 95 88 9 Pr es en t s tu dy R. so ni ni S pa ss ka ya e t S pa ss ki i, 19 71 - D en dr oc op os sy ri ac us Pi ci da e Bu lg ar ia S ofi a - EU 66 54 62 EU 66 54 90 Li ttl ew oo d et a l. 20 08 R. tu ne te ns is J oy eu x et H ou de m er , 1 92 8 - Le pt ot ila v er re au xi Co lu m bi da e Co sta R ic a G ua na ca ste - EU 66 54 59 EU 66 54 87 Li ttl ew oo d et a l. 20 08 Ra ill ie tin a sp . - C ra x ru br a Cr ac id ae Co sta R ic a G ua na ca ste - EU 66 54 58 EU 66 54 86 Li ttl ew oo d et a l. 20 08 Ra ill ie tin a sp . - Le uc on ot op ic us v ill os us Pi ci da e U SA N eb ra sk a - EU 66 54 60 EU 66 54 88 Li ttl ew oo d et a l. 20 08 Ra ill ie tin a sp . - L. v ill os us Pi ci da e U SA N eb ra sk a - EU 66 54 61 EU 66 54 89 Li ttl ew oo d et a l. 20 08 Ra ill ie tin a sp . - M an is p en ta da ct yl a M an id ae Ch in a G ua ng do ng - - O L5 97 54 0 Tu li et a l. 20 22 Ra ill ie tin a sp . - O to ty lo m ys p hy llo tis Cr ic et id ae M ex ic o Q ui nt an a Ro o CN H E 11 96 0- 71 O R2 71 64 5 - Pa nt i-M ay e t a l. 20 23 Ra ill ie tin a sp . - Si gm od on to lte cu s Cr ic et id ae M ex ic o Yu ca ta n CN H E 10 79 0- 74 O R2 71 65 0 - Pa nt i-M ay e t a l. 20 23 Fu hr m an ne tta m al ak ar tis M ah on , 1 95 8 - C ot ur ni x co tu rn ix Ph as ia ni da e Bu lg ar ia K ha sk ov o - EU 66 54 57 EU 66 54 85 Li ttl ew oo d et a l. 20 08 doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 4 of 11 Pa ro ni el la u ro ga lli (M od ee r, 17 90 ) BW 7 La go pu s l ag op us Ph as ia ni da e Fi nl an d - FM N H K N .4 34 42 - K P1 71 53 2 Pr es en t s tu dy Pa ro ni el la sp . A X 3 N eo to m a ci ne re a Cr ic et id ae U SA M on ta na FM N H K N .4 34 43 - K P1 71 53 0 Pr es en t s tu dy C al os ta ur us m ac ro pu s ( O rtl ep p, 1 92 2) B Z 1 Th yl og al e st ig m at ic a M ac ro po di da e A us tra lia Q ue en sla nd SA M 2 39 48 O R8 05 58 2 - Pr es en t s tu dy C . t hy lo ga le B ev er id ge , 1 97 5 BY 9 Th yl og al e bi lla rd ie ri i M ac ro po di da e A us tra lia Ta sm an ia SA M 9 59 2 K P1 71 52 4 - Pr es en t s tu dy Cl ad e C Ra ill ie tin a au st ra lis (K ra bb e, 1 86 9) - D ro m ai us n ov ae ho lla nd ia e Ca su ar iid ae A us tra lia Vi ct or ia - A F2 86 91 4 EU 66 54 84 O lso n et a l. 20 01 , Li ttl ew oo d et a l. 20 08 R. fu hr m an ni (S ou th w el l, 19 22 ) EV 9 C ol um ba li vi a Co lu m bi da e Th ai la nd U do n Th an i FM N H K N .4 34 35 O R8 05 59 6 O R7 95 89 1 Pr es en t s tu dy R. tr ap ez oi de s ( Ja ni ck i, 19 04 ) EC 6 Rh ab do m ys p um ili o M ur id ae So ut h A fri ca D ro nfi el d - O R8 05 58 8 - Pr es en t s tu dy R. tr ap ez oi de s ED 0 R. p um ili o M ur id ae So ut h A fri ca D ro nfi el d FM N H K N .4 34 36 , K N .3 65 0 O R8 05 58 7 O R7 95 87 9 Pr es en t s tu dy R. tr ap ez oi de s EF 4 M ic ae la m ys n am aq ue ns is M ur id ae So ut h A fri ca Be th ul ie - - O R7 95 88 0 Pr es en t s tu dy O th er sp ec ie s C ot ug ni a sp . EV 7 C ol um ba li vi a Co lu m bi da e Th ai la nd U do n Th an i FM N H K N .4 34 45 - O R7 95 89 0 Pr es en t s tu dy Sk rj ab in ia c es tic ill us (M ol in , 1 85 8) BW 8 Te tr ao u ro ga llu s Ph as ia ni da e Fi nl an d K or pi la ht i - - O R7 95 87 2 Pr es en t s tu dy S. c es tic ill us BW 9 T. u ro ga llu s Ph as ia ni da e Fi nl an d K or pi la ht i - - O R7 95 87 3 Pr es en t s tu dy 1 D M Z , d em il it ar iz ed z on e be tw ee n S ou th a nd N or th K or ea . Ta bl e 1. c on tin ue d 28S and nad1, respectively. Successfully amplified DNA was pu- rified using E.Z.N.A. TM Cycle Pure Kit (OMEGABio-Tek). Puri- fied PCR products were directly sequenced using dye terminators and visualised with an ABI 3730xl DNA analyser. Assembled sequences were submitted to GenBank (Table 1). Sequences were aligned with Muscle (Edgar 2004); ambig- uously aligned sites and gaps were deleted. The best substi- tution models, selected by the Bayesian information criterion implemented in MEGA11 (Tamura et al. 2021), were HKY+G, GTR+G and GTR+G+I for 28S, nad1 and their concatenated data, respectively. The Bayesian phylogeny inference (Huelsenbeck et al. 2001) and the Maximum Likelihood (ML) method were used for assess- ing phylogenetic relationships among the isolates. The Bayesi- an analysis was performed using MrBayes v. 3.1 (Ronquist and Huelsenbeck 2003) implemented in Geneious Pro v. 5.3.6. (http:// www.geneious.com). MrBayes was run for 5 million generations, sampled every 1,000 generations, and 500,000 generations were discarded as burn-in. Node support was expressed as posterior probabilities, values > 0.95 being considered significant. Boot- strap support for the ML method implemented in MEGA11 was based on 1,000 replications, with values > 0.75 considered sig- nificant. Three separate phylogenetic analyses were performed, i.e. for 28S, nad1 and their concatenated data. The lengths of the final alignments for these data sets were 674 bp, 708 bp and 1,885 bp, respectively. Due to amplification problems, especially for the 28S primers, the composition of the sequence data sets for the two markers is not identical. In addition to the present material, 28S and nad1 sequences for ten species of davaineids from birds and mammals were retrieved from GenBank (Table 1). The monophyly and phylogenetic relationships of davaineids among cyclophyllidean cestodes were tested using the 28S se- quences listed in Table 1, with additional sequences retrieved from GenBank, representing the families Anoplocephalidae sen- su stricto, Hymenolepididae, Dilepididae, Catenotaeniidae and Linstowiidae (Fig. 1). Mesocestoides sp. and Tetrabothrius for- steri (Krefft, 1871) were used as outgroup species for the 28S analysis, and Hymenolepis diminuta (Rudolphi, 1819) and An- drya rhopalocephala (Riehm, 1881) for the nad1 and concatenat- ed (28S + nad1) analyses. Sequences of outgroup species were obtained from GenBank. RESULTS The 28S phylogeny (Fig. 1) showed that the davaineids form a strongly supported clade among cyclophyllidean cestodes, but their relationships with other families re- mained unresolved. The Davaineidae, including Inermicap- sifer spp., is clearly not phylogenetically related with the Anoplocephalidae sensu stricto, the latter family forming a strongly supported clade with the Hymenolepididae and Dilepididae. The position of the Linstowiidae (Mathevo- taenia sp.) supports the validity and independence of the latter family with respect to other cyclophyllidean fami- lies, including the Anoplocephalidae sensu stricto These patterns are largely consistent with those of Waeschenbach and Littlewood (2017). The nad1 phylogeny (Fig. 2) showed that Skrjabinia cesticillus (Molin, 1858) is sister to the the “main clade” doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 5 of 11 Fig. 1. Phylogenetic relationships between the cyclophyllidean families Davaineidae, Anoplocephalidae sensu stricto, Hymenolepidi- dae, Dilepididae and Linstowiidae, and between isolates of the family Davaineidae (including Inermicapsifer spp.) based on sequences of the large subunit ribosomal RNA gene (28S). The topology is based on Bayesian phylogeny inference. doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 6 of 11 of the subfamily Davaineinae. Cotugnia sp. did not, how- ever, group with other davaineids (Fig. 2). Cotugnia has previously been assigned either to a separate subfamily, the Cotugniinae (see Movsesyan 2003b), or to the subfamily Davaineinae (see Jones and Bray 1994); the present result supports the former view. The phylogenies based on the three sequence data sets (Figs. 1–3) all revealed a presence of three supported line- ages (A–C) within the main clade of davaineines, although the support for lineage B was relatively low (Bayesian pos- terior probability, 0.86) in the 28S data (Fig. 1). The rela- tionships between lineages A–C remained poorly resolved. Lineage A consisted almost entirely of parasites of ro- dents, primarily of the subfamily Murinae (family Muri- dae) from South-East Asia, South Africa and Cape Verde in the Atlantic Ocean, but also endemic Malagasy rodents (Nesomys spp.) of the family Nesomyidae and Laonastes aenigmamus Jenkins, Kilpatrick, Robinson et Timmins of the family Diatomyidae from Laos (Table 1). Lineage A included four sublineages, two of which consisted of Raillietina spp. from South-East Asia, one of Raillietina spp. from Madagascar and one of Inermicapsifer spp. from South Africa and Cape Verde (Figs. 1–3). One of the South- East Asian lineages (“A1”) and the Inermicapsifer clade appeared as well supported sister lineages, especially in the concatenated data (Fig. 3). The Inermicapsifer clade also included an isolate from the pangolin Smutsia temminckii (Smuts) (Maniidae) from South Africa (Fig. 2), which is probably Inermicapsifer rhodesiensis Mettrick, 1959, de- scribed from the same host species from the present-day Zimbabwe (Mettrick 1959). The structure of the phyloge- netic trees suggests that within clade A there are at least ten independent species of Raillietina and possibly five in- dependent species of Inermicapsifer (Table 1). Lineage A also included a species of Delamuretta which appeared as a basal sublineage and is possibly sister to the clade formed by the rest of species within this lineage. Lineage B was dominated by species from birds, but it also included five species from a diverse assemblage of mammals. The analysis based on the concatenated data (Fig. 3) suggested that clade B is split in two main sub- lineages, one of which includes two species from birds (i.e. Raillietina tunetensis Joyeux et Houdemer, 1928 and Raillietina sp.) and the other includes the rest of the species within lineage B. The bird cestodes within lineage B con- sisted of several species of Raillietina and also Paroniel- la urogalli (Modeer, 1790) and F. malakartis. Lineage B also included the type species R. tetragona from galliform birds, but its phylogenetic position remained undefined, except that it showed a slightly supported relationship with Raillietina echinobothrida (Megnin, 1880) in the nad1 data (Fig. 2) and a strongly supported relationship with F. malakartis in the concatenated data (Fig. 3). The cestodes from mammalian hosts in lineage B in- cluded Raillietina coreensis Honda, 1939 from a murine rodent (Apodemus Kaup) from Korea, two unidentified species of Raillietina from cricetid rodents (Ototylomys Merriam, Sigmodon Say et Ord) from Mexico (Fig. 1) and one from a pangolin (Manis Linnaeus) from China (Fig. 2), Paroniella sp. from a cricetid rodent (Neotoma Say et Ord) from North America (Fig. 2) and two species of Calostau- rus Sandars, 1957 from Australian marsupials (Diproto- dontia) (Fig. 1). The small, strongly supported lineage C consisted of two species of Raillietina from birds, i.e. Raillietina aus- tralis (Krabbe, 1869) from Dromaius Vieillot (emu) from Australia and Raillietina fuhrmanni (Southwell, 1922) from Columba Linnaeus from Thailand, and Raillietina trapezoides (Janicki, 1904) from South African murine ro- dents, the latter two species showing a strongly supported relationship (Figs. 1–3). DISCUSSION Phylogenetic patterns The main phylogenetic pattern emerging from the pres- ent analysis is the presence of three independent lineages within the main clade of the subfamily Davaineinae, one of which (A) is almost entirely confined to species from rodents and the other two (B, C) show a mixture of species from birds and mammals. Based on the facts that lineages B and C evidently originated in birds and that the basal genera Cotugnia and Skrjabinia are parasites of the same host group, the most parsimonius assumption is that the original hosts of the main clade of the Davaineinae are found among birds. The species in the genus Skrjabinia, the sister group of the main davaineine clade, are predom- inantly parasites of galliform birds (Movsesyan 2003a). Further, the presence of five species of cestodes from gall- iforms in lineage B (Fig. 2) leads us to suggest that this group of birds harboured the tapeworm lineage that gave rise to the main davaineine clade. However, the basal spe- cies in lineage C (Raillietina australis) is a parasite of ratite birds (Dromaius), which appeared before galliform birds. Therefore, we cannot rule out the possibility that the ori- gin of the Davaineinae is with the ratites. The Australian emu, Dromaius novaehollandiae (Latham), has at least five host-specific species of Raillietina (see O’Callaghan et al. 2000). The three main lineages, particularly those dominated by parasites of birds (B, C), showed very extensive geo- graphic distributions. For example, the range of clade B covers Europe, China, Korea, South-East Asia, North America and Central America (Fig. 2), and possibly also Australia and Mexico (Fig. 1). It is probable that the high mobility of birds, including their migratory behaviour, is the main factor explaining the observed pattern. The major diversification of the precursor of the main clade of davaineines into three lineages (A–C) remains partly obscure, but it is clear that there was an early colo- nisation of rodents (lineage A) from birds, probably gall- iforms. The species in lineage A are almost exclusively parasites of murine rodents (Figs. 1–3). In the nad1 and concatenated trees, the basal relationships within lineages A were unresolved, but the structure of the 28S tree (Fig. 1) suggests that there is a basal South-East Asian species (isolate CI1), which is sister to the clade formed by the rest of the species in the lineage A. The presence of sev- doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 7 of 11 Fig. 2. Phylogenetic relationships between isolates of the family Davaineidae (including Inermicapsifer spp.) based on sequences of the mitochondrial NADH dehydrogenase subunit 1 gene (nad1). The topology is based on Bayesian phylogeny inference. eral additional isolates from South-East Asia in the main sublineage suggests that lineage A originated in South-East Asian murines. Sublineage A1 probably also originated in South-East Asian murines, with a subsequent colonisa- tion of African murines, leading to divergence and diver- sification of the Inermicapsifer clade. These patterns are consistent with the phylogenetic history for the subfamily Murinae, which probably appeared in South-East Asia and subsequently colonised Africa and other regions (Schenk et al. 2013). The present study includes the first report of davaineid cestodes in endemic Malagasy nesomyid rodents (Nesomys spp.) of the subfamily Nesomyinae. The phylogenetic po- sition of these cestodes indicates an early divergence with- in lineage A. The Malagasy nesomyid rodents probably originate from a single colonisation event by an African nesomyid 24–20 Mya (Poux et al. 2005), suggesting that their extant, host-specific parasites have an equally long history on Madagascar. However, a species of Raillietina (R. murium Joyeux et Baer, 1936) has been described from doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 8 of 11 the black rat Rattus rattus from Madagascar (Joyeux and Baer 1936), which leads to an alternative hypothesis that the Raillietina species of the endemic nesomyids originate through a recent capture from commensal rats. However, the latter hypothesis is invalid owing to the basal phyloge- netic position and high genetic divergence of the Raillieti- na species of nesomyids within lineage A. A recent survey of helminths of Malagasy rodents showed an absence of davaineid cestodes in black rats (n = 90) and Eliurus spp. (Nesomyidae) (n = 17) (JTL and VH – unpubl. data). There are evidently no reports of Raillietina from African ne- somyids (subfamilies Cricetomyinae, Delanymyinae and Dendromurinae). Laonastes aenigmamus or the Laotian rock rat (“kha- nyou”) is a recently found “enigmatic” rodent that was shown to belong to the Diatomyidae, a fossil family thought to be extinct for 11 My (Dawson et al. 2006). It has a limited distribution in the karst region of Central Laos (Nicolas et al. 2012) and also in a small, adjacent region of Vietnam (Nguyen et al. 2014). Hugot et al. (2013) de- scribed a new species and genus of an oxyurid nematode from L. aenigmamus, but there are no published reports of its tapeworms. The structure of the present phylogenetic trees suggests strongly that the Raillietina species of L. ae- nigmamus originate through a single host shift from South- East Asian murine rodents (Figs. 1–3), and are therefore much more recent than the host itself. Lineage B probably diversified initially in galliforms, with subsequent shifts to woodpeckers (Piciformes), doves (Columbiformes) and mammals. The three Raillietina spe- cies from woodpeckers appeared as a monophyletic group (Fig. 2), as shown earlier by Littlewood et al. (2008). Among the mammalian hosts, there were at least two tape- worm species from rodents and one from pangolins (Fig. 2), which evidently differentiated as a consequence of a shift from birds. The 28S tree suggests that clade B also includes two additional species from cricetid rodents (Raillietina spp.) from Mexico and two from marsupials (Calostaurus spp.) (all without nad1 data), although the support for clade B was relatively low (Fig. 1). Additional phylogenetic analyses including unpublished 28S and nad1 sequenc- es of Raillietina sp. from a “field biologist” (KT001065, KT001066; submitted to GenBank by B. A. Kendall, V. V. Tkach et al.) showed that the species in question is a parasite of birds within clade B (results not shown). This finding attests to the ability of bird-associated davaineids to infect mammalian hosts. The original bird hosts for clade C remain unknown, but it is obvious that murine rodents were colonised by a tape- worm lineage of birds, probably columbiforms, giving rise to (the precursor of) R. trapezoides (Figs. 1–3). Based on the present results, there has been in total 5–7 independent colonisations of mammals by davaineine spe- cies of birds, and two colonisations by parasites of murine rodents to other mammals (a diatomyid rodent, a pango- lin). No shifts from mammals to birds were inferred. Al- though no explicit cophylogenetic analysis was performed, it is clear that the colonisation of mammals by parasites of birds and other mammals has played a major role in the diversification of the main davaineine clade. No unambig- uous evidence for cophylogenetic events of the host and parasite lineages was found. Systematics The present phylogenetic analyses showed consistently that all isolates of Inermicapsifer form a strongly support- ed monophyletic group, which is positioned among Railli- etina spp. in the “rodent lineage” (A). Therefore, the genus Inermicapsifer is unambiguously a member of the family Davaineidae, specifically the subfamily Davaineinae, and should be excluded from the family Anoplocephalidae. The implication is that the inermicapsiferines are davain- eids which have lost the rostellum and armature in con- nection with their divergence from other davaineids of ro- dents (Raillietina spp.) and that the loss has occurred only once. This also means that the subfamily Inermicapsiferi- nae and the family Inermicapsiferidae should be treated as synonyms of the Davaineidae, specifically the subfamily Davaineinae. The independence of inermicapsiferines with respect to the Anoplocephalidae sensu stricto is supported by ultrastructural characteristics of spermatozoa (Miquel et al. 2016) and eggs (Świderski et al. 2015a). The species of the genus Inermicapsifer are primarily parasites of hyraxes (Hyracoidea), with the type species In- ermicapsifer hyracis (Rudolphi, 1808), and rodents, mainly in sub-Saharan Africa (see Caira et al. 2024). Morphologic evidence concerning the structure of the osmoregulatory system and position of the female glands suggests that the Inermicapsifer species of rodents represent a separate, new genus. However, the present analysis lacks the inermica- pasiferines of hyracoids, and without knowledge of their phylogenetic position, an erection of a new genus would be premature. In addition to Inermicapsfer, the genera traditionally assigned to the Inermicapsiferinae include Metacapsifer Spasskii, 1951, Pericapsifer Spasskii, 1951 and Thysano- taenia Beddard, 1911; Arhynchotaenia Pagenstecher, 1877 and Hyracotaenia Beddard, 1912 are junior synonyms of Inermicapsfer (see Spasskii 1951 and Caira et al. 2024). Metacapsifer, Pericapsifer and Thysanotaenia, which share the main morphological features with Inermicapsfer, including egg-bearing parenchymtous capsules, are here assigned to the Davaineidae (subfamily Davaineinae). The fact that Raillietina spp. are present in all three main lineages (A–C) and appear as multiple independent sublin- eages from bird and mammalian hosts unambiguously ver- ifies the non-monophyly of the genus Raillietina. Most of the Raillietina lineages identified here evidently represent new genera. The question which species should be includ- ed in Raillietina sensu stricto cannot be reliably answered based on the present results, because the phylogenetic re- lationships among the species in lineage B, including the type species R. tetragona, remained largely unresolved. However, the phylogenetic association between R. tetrag- ona and R. echinobothrida, both from galliform birds, was strongly supported by Siddiqui et al. (2023), and slightly supported by Butboonchoo et al. (2016) and by the present doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 9 of 11 Fig. 3. Phylogenetic relationships between isolates of the family Davaineidae (including Inermicapsifer spp.) based on concatenated sequences of the mitochondrial NADH dehydrogenase subunit 1 gene (nad1) and the large subunit ribosomal RNA gene (28S). The topology is based on Bayesian phylogeny inference. nad1 analysis (Fig. 2). The genus Raillietina is obviously in need of an extensive reorganisation and description of new taxa based on phylogenetic and morphologic criteria. The present results also provide evidence for non-mono- phyly of the genus Paroniella. Acknowledgements. For the studies conducted in South Africa, private landowners and reserve managers are thanked for access to the properties. Rodents were sampled under the following pro- vincial permit numbers: Eastern Cape, CRO37/11CR; Free State, 01/8091 and Gauteng, CPF 6–0153. All applicable institutional and/or national guidelines for the use of animals were adhered to reference numbers: 2006B01007 and SU-ACUM11–00004 (Stel- lenbosch University); AEC32.11 (Agricultural Research Institute – Onderstepoort Veterinary Institute). Several researchers pro- vided field and technical support. This work was financially sup- ported by the National Research Foundation (NRF), Agricultural Research Council–Onderstepoort Veterinary Institute and Stel- lenbosch University. The Grant holder acknowledges that opin- ions, findings and conclusions or recommendations expressed in any publication generated by the NRF-supported research are those of the authors, and that the NRF accepts no liability what so ever in this regard. The studies performed in South-East Asia were funded by the French National Research Agency (ANR), project CERoPath (ANR 07 BDIV 012). The collection of hel- minths from Cape Verde was supported by the Spanish Ministry of Foreign Affairs and Cooperation (A1/035356/11). Ian Bever- idge and Marja Isomursu are acknowledged for kindly supplying tapeworms from Australia and Finland, respectively. Two review- ers provided helpful comments on the manuscript. doi: 10.14411/fp.2024.011 Haukisalmi et al.: Phylogenetics of the Davaineidae (Cestoda) Folia Parasitologica 2024, 71: 011 Page 10 of 11 Author contribution statement. The first author is responsible for the design of the study, identification of the parasites, molecular and phylogenetic analyses, and writing of the manuscript, The other au- thors designed and performed the collection of the host and parasite specimens, and contributed to the writing of the manuscript. REFERENCES Baer J.G., Fain A. 1955: Les Cestodes des Pangolins. Bull. Soc. Neuchâtel. Sci. Nat. 78: 37–52. Beveridge I. 1994: Family Anoplocephalidae Cholodkovsky, 1902. In: L.F. Khalil, A. Jones and R.A. Bray (Eds.), Keys to the Cestode Parasites of Vertebrates. CABI, Wallingford, pp. 315–366. 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Cite this article as: Haukisalmi V., Ribas A., Hugot J.-P., Morand S., Chaisiri K., Junker K., Matthee S., Spickett A., Lehtonen J.T., Feliu C., Henttonen H. 2024: Phylogenetic relationships and systematics of tapeworms of the family Davaineidae (Cestoda: Cyclophyllidea), with emphasis on species in rodents. Folia Parasitol. 71: 011. Received 12 March 2024 Accepted 24 April 2024 Published online 6 June 2024