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Polyphyly and Monophyly in the Family Linstowiidae : Cestode (Tapeworm) Parasites of Reptiles, Birds, Monotremes, Marsupials and Placentals Mammals. Mariel Campbell Museum of Southwestern Biology, UNM Fall 2018. Linstowia spp.
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Polyphyly and Monophyly in the Family Linstowiidae: Cestode (Tapeworm) Parasites of Reptiles, Birds, Monotremes, Marsupials and Placentals Mammals Mariel Campbell Museum of Southwestern Biology, UNM Fall 2018
Marsupials evolved in Asia and North America, radiated in South America, and crossed Antarctica, to Australia 30-50 million years ago (Nilsson et al 2010). Did they take their tapeworms with them?
Question 1: Does the Genus Linstowia represent a monophyletic group? • Hypothesis A: The genus Linstowiais monophyletic, with South American and Australian clades forming sister taxa. South American and Australian Linstowia species derive from a shared ancestor prior to the separation of the continents. • Hypothesis B: • The genus Linstowia is polyphyletic or paraphyletic. South American and Australian species of Linstowia are more closely related to other linstowiid taxa than to each other. Similarities between South American and Australian species of Linstowiaare due to parallel evolution and/or independent acquisition from reptile ancestors.
Morphological character matrix of 69 characters (28 multistate, 40 binary) for 26 Linstowiid taxa All multistate characters treated as unordered using Fitch parsimony. Taxa with more than one character state were interpreted as polymorphic. All characters were rescaled to give binary and multistate characters equal weight. Trees were initially rooted with midpoint rooting to verify that ingroup taxa were monophyletic, subsequently, outgroup rooting was used with two species of Oochoristica from reptiles.
Linstowiais not monophyletic based on this analysis of morphological characters. Maximum Parsimony Analysis Results PAUP 4.0 26 taxa 68 morphological characters scaled for equal weights, base weight 1000 Heuristic Search, TBR, with 1000 Bootstrap replicates : 1 Tree Length: 186078 CI: 0.36 RI: 0.58 RC: 0.21 HI: .64 “Linstowia”= Neotropical Marsupials OochoristicaeremophilaAustralia Atriotaenia New World skunks, raccoons Paratriotaenia New World primates OochoristicaantechiniAustralia Mathevotaenia spp. New world marsupials and edentates “Linstowia” =Australian Marsupials and Monotremes Oochoristica : New World and Australian reptiles
Question 2: What is the status of the Family Linstowiidae? • Despite elevation to family by Spasskii 1951, the “Family” Linstowiidae has not been recognized in the literature. • Placed historically in the Family Anoplocephalidae. • Are Linstowiid cestodes a subfamily of Anoplocephalidae or a separate family? • E.g. Do they share a most recent common ancestor with anoplocephalids or with other taxa?
Relationship of Linstowid cestodes to Family Anoplocephalidae and other Families in Order Cyclophyllidea: • Methods: • 1) Search GenBank: single linstowid sequence (Mathevotaeniasymmetrica, 28S) • Blasted sequence against other Cestoda • Selected representatives of other families in Order Cyclophyllidea with 28S sequence data; • Choose representatives with lowest E-scores and greatest sequence overlap • Selected outgroup taxa in Order Diphyllobothridea • Downloaded and aligned sequences of 46 cestode taxa • Compared alignments by Muscle and Clustal Omega • Trimmed and untrimmed sequences • 5) Maximum Parsimony analysis in Paup 4.0 • RaxML analysis • Bootstrapped 1000 replicates • Analyzed resulting trees
PAUP Maximum Parsimony Analysis of 46 Cestode Taxa Using 28S nuclear rRNA Outgroup rooting Trimmed alignment 1000 BS replicates Heuristic Search Default settings Tree Length 6008 CI RI RC HI 0.54 0.66 0.36 0.46 Anoplocephalidae Linstowiidae
-27419.116740 RaxML Analysis of 46 Cyclophyllidean Cestode Genera using nuclear ribosomal 28S 100 Bootstrap replicates GTR Gamma+I TBR Heuristic 1 Tree -ln L = 29071 27419.116740 AIC = 58329
Summary • Morphological analysis suggests that the genus Linstowia does not represent a monophyletic group sharing a common ancestor dating from prior to the separation of the continents. • Based on this analysis, the genus represents separate South American and Australian lineages that should be split into separate genera. These lineages may have been acquired separately from parasites of reptiles. • Based on molecular analysis of two sequences, Linstowiid cestodes are evolutionarily distinct and should be a separate family distinct from cestodes in the Family Anoplocephalidae. • This study provides evidence that carrion beetles may be a potential intermediate host for the Family Linstowiidae. • The group is very poorly known with many interesting, ancient, and potentially informative lineages that can be examined to study host and parasite evolutionary and phylogeographic patterns. But we need molecular samples.
References Beveridge, I. "The genus LinstowiaZschokke, 1899 (Cestoda: Anoplocephalidae) in Australian mammals with the description of a new species, L. macrouri." Systematic Parasitology 5.4 (1983): 291-304. Caira, Janine N., and Kirsten Jensen. Planetary biodiversity inventory (2008–2017): Tapeworms from vertebrate bowels of the earth. Natural History Museum, University of Kansas, 2017. Janicki, C, . VON. 1906. Studien an Siugetiercestoden. Zeitschrift fir Wissenschaftliche Zoologie 81: 505-597. Kaltenpoth, Martin, and Sandra Steiger. "Unearthing carrion beetles' microbiome: characterization of bacterial and fungal hindgut communities across the S ilphidae." Molecular Ecology 23.6 (2014): 1251-1267. Nilsson MA, Churakov G, Sommer M, Tran NV, Zemann A, et al. (2010) Tracking Marsupial Evolution Using Archaic Genomic Retroposon Insertions. PLOS Biology 8(7): e1000436. https://doi.org/10.1371/journal.pbio.1000436 Pais, F. S., Ruy, P. C., Oliveira, G., & Coimbra, R. S. (2014). Assessing the efficiency of multiple sequence alignment programs. Algorithms for molecular biology : AMB, 9(1), 4. doi:10.1186/1748-7188-9-4 Schiewe, J. (28 July 2010). "Australia's marsupials originated in what is now South America, study says". Los Angeles Times. Archived from the original on 1 August 2010. Retrieved 1 August 2010. Schmidt, G, . D. 1986. Handbook of tapeworm identification. CRC Press, Boca Raton, Florida6 Spasskiĭ, AlekseĭAndreevich. Anoplocephalate tapeworms of domestic and wild animals. Vol. 1. Academy of Sciences of the USSR, 1951 ie Jerusalem, Israel Program for Scientific Translations, 1951. Wiens, John J. "Character analysis in morphological phylogenetics: problems and solutions." Systematic Biology50.5 (2001): 689-699. Zschokke, F., 1899. Neue Studien an Cestoden aplacentaler Saugethiere. Zeitschrift fur WissenschaftlicheZoologie 65: 404-445. Zschokke, F. 1904a. Die Darmcestoden der amerikanischen Beuteltiere. Centralblatt fur Bakteriologie, Parasitenkunde und Infektionskrankheiten, 1st Abt. 36: 51-62. Zschokke, F. 1904b. 2. Die Cestoden der siidamerikanischen Beuteltiere. Zoologischer Anzeiger 27: 290- 293.
Acknowledgments: Dr. Scott L. Gardner Bolivia Field Crews 1983-1993