A checklist, including all the cestode taxa recorded from Switzerland, their hosts, as well as deposited specimens available in scientific collections, is provided. The country has one of the richest European cestode faunas consisting of 251 species, almost all of them cyclophyllideans, that were identified in 190 vertebrate and 24 invertebrate host species. This is a very significant increase over the previous similar list that was established one century ago by Fuhrmann (1926). Since then, advances have been particularly important for parasites of mammals and birds although an important margin of progress remains for the latter as several bird families have been surprisingly little studied in the country. A large proportion of species described in Switzerland, including 22 that are represented by types, are available in public collections, most of them at the Muséum d'histoire naturelle de Genève. New reports were numerous in the second half of last century but have become scarce in recent decades. Today, tapeworms have been identified in no more than one third of Swiss vertebrate species and despite one century of progress, the true diversity of this fauna in the country remains to be determined.
INTRODUCTION
Human-created borders obviously do not limit biological taxa, and it may look somewhat archaic to establish national checklists for the latter. However, despite their artificiality, such lists remain useful as practical tools in many domains, not least for conservation management. Furthermore, they allow aggregating hard to find and dispersed data, especially for poorly studied groups; or facilitate faunistic comparisons between regions. Although checklists are regularly published for better known groups, like vertebrates or some arthropods, this is generally not the case for less popular taxa, whatever their diversity or ubiquity. Parasitic helminths, and among them cestodes, certainly belong to this category, even though Switzerland was home to some of the most famous and prolific cestodologists of the 20th century, like Otto Fuhrmann (1871-1945) or Jean-Georges Baer (1902-1975). However, although these authors, and their students, contributed significantly to knowledge of the Swiss fauna, they never particularly focused on it. Today, the single exhaustive list of Cestoda found in Switzerland remains that of Fuhrmann (1926) published almost a century ago.
Similar global checklists are also rare for most other countries, the most notable and recent exception in Europe being the “Checklist of tapeworms of vertebrates in Finland” (Haukisalmi, 2015). A few other ones in Belarus (Merkusheva & Bobkova, 1981), the Iberian Peninsula (Cordero del Campillo et al., 1994), Slovakia (Macko et al., 1993, 1994; Hanzelova et al., 1995; Hanzelova & Ryšavý, 1996, 1999) and Poland (Pojmanska et al., 2007) also exist. An ancient checklist for France (Joyeux & Baer, 1936) was ill named, as it also covered many taxa absent from this country. On-line checklists are available for Italy ( http://www.faunaitalia.it/checklist/index.html) (Stoch, 2003), as well as for the United Kingdom ( https://www.nhm.ac.uk/research-curation/scientific-resources/taxonomy-systematics/host-parasites/database/index.jsp) (Natural History Museum, London, 2007), although with limited updates or accompanying information. A recent and comprehensive list for Western Europe is therefore lacking. The goal of this study is to provide a complete summary of the known fauna of cestodes in Switzerland. This list includes the names of the parasites, their known hosts, a simplified distribution in the country, a selective bibliography as well as a list of the known specimens kept in scientific institutions.
It should be emphasized that such a compilation is not a taxonomic work. As far as possible, historic names of parasites are listed under their presently accepted synonymy and they are placed in the most recent higher systematics of the group (Caira & Jensen, 2017), however no nomenclatural act is made here. This checklist aims at being a practical reference and tool for researchers and other users of biodiversity information.
HISTORICAL CONTEXT
The earliest report of an identifiable parasitic worm in Switzerland seems to be that of Diphyllobothrium latum by Dunus (1592). It was followed by a few publications in the 17th century on parasites of humans. Later reports became more common but remained mostly limited to common parasites of domesticated animals until the mid-19th century when a more diverse and steady flux of publications started. A rather large spectrum of authors has contributed to the knowledge of the Swiss fauna, but only a few have built a consistent body of work. The most important of them being Friedrich Zschokke (1860-1936) in Basel, who published essentially on parasites of fish; Bruno Galli-Valerio (1867-1943) in Lausanne, veterinarian and physician, who probably examined the largest diversity of Swiss vertebrates through numerous publications (see Gaschen, 1950); Otto Fuhrmann (1871-1945) in Neuchâtel, one of the giants of cestodology, who worked on most tapeworm groups and contributed significantly to the knowledge of Swiss Cyclophyllidea; Bernd Hörning (1931-2012) in Bern, a veterinarian with interest in many wild animals (see in particular: Hörning, 1963); and Claude Vaucher (1942-) in Geneva, who decisively contributed to the knowledge of the helminthofauna of micromammals.
Besides parasites of domestic animals (and humans), the focus of cestodes biodiversity research in Switzerland long concentrated on parasites of fish from the country's major lakes, resulting in plethoric, and often repetitive, observations, especially at the turn of the 20th century. These are particularly problematic when considering the extremely complex fish diversity in Alpine lakes (see below). Works by K. Wolffhügel (1869-1951) or Fuhrmann then progressively expanded our knowledge of the bird fauna, but a new focus on wild mammals only appeared in the 1960s. However, the latter two classes of vertebrates remain relatively poorly known, with significant gaps for example in passerines, or in chiropterans. As for amphibians and reptiles, only a very few isolated publications exist. Reports from larval forms in invertebrates are also limited and mostly ancient, as research on life cycles progressively became unfashionable. The last significant reports in this field in Switzerland are from the late 1980s (Szelenbaum-Cielecka et al., 1988).
The last two new cestode species descriptions from Swiss vertebrates, one in very common Song Thrushes and one in Great crested Grebes, are respectively over 30 and 20 years old (Gigon & Beuret, 1991; Vasileva et al., 2000).
MATERIAL AND METHODS
Sources:
1) Museum data. Muséum d'histoire naturelle de Genève (MHNG) holds one of the major collections of cestodes worldwide (>22'000 lots), including most historic collections of Swiss cestodologists. Its catalogue provided the backbone of the present list. Additional data were requested from other Swiss Museums that maintain scientific collections as well as from major foreign Museums likely keeping Swiss specimens.
2) Other institutions data. Some veterinary or paramedical institutions in Switzerland were asked for archives or registers.
3) Bibliography. Searches were made with appropriate taxonomical and geographical filters in Web of Science's Zoological Record (© Clarivate).
4) Several unpublished student works from parasitology departments in Swiss institutes.
Conditions for listing:
For a species to be included in the checklist, at least one of two nonexclusive conditions had to be met: The taxon had to be published with an explicit mention of its Swiss origin and/or specimens labeled as of Swiss origin had to be registered in collections in an academic institution. For the sake of consistency, and contrary to Fuhrmann (1926), observations from bordering foreign localities (e.g., “Salève” or “Black Forest”) are not included here, leading to the exclusion of a few taxa that were listed in Fuhrmann's catalogue.
Data provided (Table 1, Annex 1):
Cestodes specific identification (generic in a few cases): Except in rare instances, identifications have not been checked. Thus, the original publication/label name is reported, either as the valid or synonym name of the taxon. A synonym list is provided but does not aim at comprehensiveness: only names that have been used when specifically referring to specimens in Switzerland are mentioned.
Hosts: All hosts, both final and intermediate, belonging to the Swiss fauna are reported.
For vertebrates, hosts are listed under their present taxonomic status according to Fishbase (Froese & Pauly, 2021), Amphibian Species of the World (Frost, 2021), The Reptile Database (Uetz & Etzold, 1996), Avibase (Lepage, 2018), and Aulagnier et al. (2008) respectively for fish, amphibians, reptiles, birds and mammals (all online references accessed in 2021). Ancient host names have been updated in accordance. Parasites of captive hosts not belonging to the Swiss fauna are not listed.
The Swiss vertebrate fauna is well known, and most names are unambiguous. A significant exception to this statement concerns the fish, and especially the whitefishes (Coregonus spp., Salmonidae). Over the years, a very large number of names, both scientific and vernacular, have been used for these fish, especially in Swiss lakes. This is due to a complex history of speciation, colonization, hybridization, human transfers, and local extinctions. Revisions and descriptions of new taxa have been numerous (e.g., Selz et al., 2020). Up to 35 species may presently be living in Swiss lakes, but revisions keep diverging both in the number and names of these taxa. Despite several attempts to clarify their systematic status (e.g., Steinmann, 1950; Kottelat & Freyhof, 2007) no taxonomical consensus presently exists. Even the species concept best adapted to Coregonus taxa is not clear as so called “speciation reversals” seem to be the norm under changing ecological conditions (Vonlanthen et al., 2012). The problem is further enhanced when trying to match often partial or imprecise historical observations to present day nomenclature, a close to impossible task. Hence, I chose to retain the limited number of Coregonus species names that are recognized as valid in Fishbase (Froese & Pauly, 2021) even though this can lead to some inconsistencies. For example, in Lake Geneva, historical species are now considered extinct (Vonlanthen et al., 2012), but I nevertheless use their names, as in the original publications, as no consensus exists on other ones. It is most likely that each significant water body in the country hosts its own fish population/parasite population fauna (with possible speciation for either or both in some cases), but no convenient nomenclatural system, neither for hosts nor for parasites, has yet been accepted to adequately represent this situation.
In addition, some rare ambiguities may occur for reports from domestic mammals that have a wild conspecific in the country, in particular for cats, with both Felis s. silvestris and F. s. catus living in Switzerland. Unless specified otherwise, reports are assumed to be from the domesticated form.
Developmental stage: Hosts of larval forms are mentioned as such. Note that in a few cases both adult and larval worms can be found in the same host.
Localities: Detailed localities are not reported (and, most often, not available); only cantons and large water bodies are mentioned when such information exists. In some cases, especially for collection specimens, only the mention “Switzerland” is available, resulting in this field being kept empty. Cantons and water bodies mentioned might in some cases refer to the same observation from different sources.
Standard Swiss cantons abbreviations ( https://www.iso.org/obp/ui/#iso:code:3166:CH) are used, except for BA (Basel) being used as a collective for BL and BS (Basel state and Basel city). The main water bodies are abbreviated as follows: A: Lake Maggiore; B: Lake Biel/ Bienne; L: Lake Geneva/Léman; M: Lake Morat; N: Lake Neuchâtel; O: Lake Constance/Bodensee; T: Lake Thun; U: Lake Zug; V: Lake Lucerne/Vierwaldstättersee; Z: Lake Zürich. In a few cases CH is used for a documented countrywide distribution.
Collection dates: Only unambiguously reported collection years (which can significantly differ from publication dates) are mentioned. In most cases, especially for more ancient records, this information is lacking.
References: Bibliographic references are not listed exhaustively. A subjective selection of the most relevant publications citing the taxon is mentioned. For many specimens in collections, no associated publication is known.
Specimens: All databased specimens I am aware of are listed here with their accession number, and type status where appropriate. There are, however, a few unregistered samples from the large common species (Taenia, Diphyllobothrium, Ligula, …) on display, or in the collections, of many smaller institutions. Museum acronyms: IPCAS: Institute of Parasitology, Czech Academy of Sciences; MHNF: Musée d'histoire naturelle, Fribourg; MHNG-PLAT: Muséum d'histoire naturelle de Genève, Platyhelminthes Collections; MUW: Department of General Biology and Parasitology, Medical University of Warsaw; GBIFCH: Musée de Zoologie, Lausanne, Invertebrates collections (=MZL-Invert); NMB-CEST: Naturhistorisches Museum, Basel, Cestodes collections; NHM: Natural History Museum, London; NWSW: Naturmuseum Winterthur; USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC; ZMZ: Zoologisches Museum Zürich.
RESULTS
Mentions of at least 251 cestode species [in 125 genera and 21 families] forming 689 host/parasites pairs could be traced in Switzerland (including 5 species inquirendae) (Tables 1, 3). This is, respectively, a 99 and 132% increase on Fuhrmann's (1926) list. The main cestode order present in Switzerland is, by far, the Cyclophyllidea (218 spp. or 87% from total) with Hymenolepididae (111 spp.) followed by Dilepididae (36 spp.) as the most represented families. Altogether 214 species of hosts, 24 invertebrates and 190 vertebrates, have been recorded harboring cestode parasites. The cestode fauna of birds is the most diversified (Table 2). Eight species are known only from their metacestodes.
Specimens from 208 (84%) of the species known from Switzerland are preserved in academic institutions, including 6 holotypes, 1 lectotype, 14 syntypes, 15 paratypes and 2 “types” specimens/lots representing 22 cestode species. These are distributed in 1250 lots, over 93% of them kept in the Muséum d'histoire naturelle de Genève (Annex 1). About 82% of the specimens with associated collection data were gathered in the second half of the last century. Parasites were recorded from all over Switzerland, although with a marked bias toward the Southern Alpine and Western parts of the country (Table 1).
REMARKS
a) Numbered remarks in Table 1:
[1] Locality uncertain and host probably Marmota sp. (Global Cestode Database, Caira et al., 2023)
[2] Possibly imported from Eastern Europe.
[3] This material is wrongly reported as D. columbina instead of D. columbae in the USNM database.
[4] As Taenia blanchardi in Fuhrmann (1926).
[5] Possibly also Galli-Valerio 1929 in VD (Gaschen, 1950).
[6] One record (MHNG-PLAT-55742) of this species is reported from Aythya marila (Anatidae), a probable mislabeling.
[7] Observation in a zoological garden, but the host is present in Switzerland.
[8] Dubious as only h. hibernia Montgomery, Montgomery & Dunn seems to parasitize Apodemus (Nkouawa et al., 2016)
[9] Marked “h. phasianina”.
[10] These records are dubious and likely due to mislabeling as these species are parasites of shrews (V. Haukisalmi, pers. communication).
[11] One slide (MHNG-PLAT-40931) is marked with Mergus serrator as host, but the specimens are misidentified.
[12] Original report mentions Anser arvensis, interpreted as A. fabalis.
[13] Published information (Vaucher & Hunkeler, 1967); however, the single matching slide in collections (MHNG-PLAT-18532) indicates R. straminea.
[14] Dubious. Vaucher (1971) considers the taxon as a specific parasite of Crocidura.
[15] Possibly also intermediate host (Eckert & Deplazes, 2004).
[16] Both in wild and domestic cats (Gaschen, 1950).
[17] Uncertain. Reported by Fuhrmann (1926) from an observation of Galli-Valerio (1916).
[18] According to Hörning (1963), only imported hares were positive.
[19] Domestic.
[20] Host inferred.
[21] These records are suspicious as rodents are normal intermediate hosts of V. mustelae.
[22] Diphyllobothrium sp. interpreted as D. latum.
[23] According to Wicht (2008), D. latum does not develop in coregonids and probably also not in salmonids. Reports in these hosts may concern D. dendriticum (or possibly D. ditremus).
[24] Local contamination but imported intermediate host.
[25] Proteocephalus host list. Multiple confusions due to the close morphological similarity between P. longicollis with P. exiguus, P. fallax, P. alosa (now synonymized), P. filicollis and P. percae have been the norm during most of the 20th century. This resulted in the mention of these worms in a variety of hosts, but many of these are most certainly misidentifications or accidental infections.
[26] P. fillicolis is a parasite of Gasterosteus aculeatus and Coregonus fera is probably a postcyclic host (i.e., an additional host becoming infected with an adult worm through predation).
[27] P. longicollis is a parasite of Salmonidae, but its presence in Alosa agone is possible. Reports in other fishes are likely misidentifications. Report in Natrix is certainly accidental/ postcyclic. M. leuckarti has been found to be an unsuitable experimental host (T. Scholz, pers. communication).
[28] A recent molecular analysis (Brabec et al., 2023) suggested, however, that P. fallax may be a valid species parasite in Coregonus sp. while P. longicollis would be restricted to Salmo spp. hosts.
[29] P. percae is a parasite of Perca fluviatilis and Esox lucius (postcyclic). Proteocephalus ocellatus (Rudolphi, 1802) was not recognized by Fuhrmann (1926) and is considered a synonym of P. percae (Muller, 1780) by Scholz & Hanzelova (1999). The numerous mentions of “P. ocellatus” in fish of other families, mostly in old records (e.g., Zschokke, 1884; Nufer, 1905) are most likely accidental or misidentifications.
[30] P. torulosus is a parasite of Cypriniforms and records in other fishes should be considered accidental or misidentifications.
b) Other remarks
Two taxa reported by Fuhrmann (1899), Acoleus vaginatus (Acoleidae) in himantopus himantopus and Gyrocoelia perversus (= perversa) (Dioicocestidae) in Limosa lapponica are not considered herein. This material was given for determination to Fuhrmann by the MHNG, but I could find no indication that it originated from Geneva (or Switzerland). As no further reports of these taxa have been published, their presence in the country remains uncertain.
Fuhrmann (1926) similarly reports the presence of Diplophallus polymorphus in Recurvirostra avosetta in Basel. A possible match for this material could be MHNG-PLAT-55673 that originates from the University of Neuchâtel collection, although no locality is mentioned on the label. Furthermore, the specimens, originally reported by Wolffhügel (1900), come from a “Zoologischer Garten von der Nordsee”. In consequence this taxon has most likely not been found in Switzerland and I haven't considered it in the table.
An occurrence of Grillotia erinaceus (van Beneden, 1858) is mentioned in Lota lota in the early literature (e.g., Zschokke, 1903; Fuhrmann, 1926). Both the freshwater host and the locality (Lake Geneva) of this single record are highly improbable for a trypanorhynch cestode. No material is known. I have removed this host-parasite occurrence from the list.
A type of Proteocephalus abcisus [= Choanoscolex abcisus (Riggenbach, 1895) La Rue, 1911] is registered from Switzerland in the USNM (#1349984). Origins of this material are unclear, but the species is from the Neotropics and does not belong to the Swiss fauna.
Some data of Vaucher (1971) are difficult to interpret as a detailed host-parasite list by locality is not given. Geographical locations were ascertained on labels linked to specimens when available. In a few cases I considered that the parasite was present in Switzerland in each of its reported hosts whose distribution encompassed the country.
DISCUSSION
Sources
It should be noted that an important part of the data collected in this work comes from natural history collections material, highlighting the crucial importance of these institutions for our understanding of the biodiversity through time. Given their highly specialized nature, only a few museums maintain scientific collections of tapeworms and therefore I assume that a very high proportion of the existing information could be accessed. A similar level of confidence could also be reached for published information through the rich bibliographic database of the MHNG library and bibliographic software. A few host/parasite reports were nevertheless difficult to track, especially when published in very local veterinary journals and a few have certainly been missed. Globally, though, I am confident that the information gathered in Table 1 is comprehensive. In addition to these traditional sources, a single occurrence of an ususual and quite unexpected host-parasite association was revealed through DNA sequencing (Taenia martis in h. sapiens, see Table 1). This is not surprising as only few sequences of cestodes of Swiss origin, mostly from Taeniidae and Proteocephalidae, are available in Genbank.
Available material
It is remarkable that a very high proportion (84%) of the species known from Switzerland are represented by at least one sample in academic institutions (Annex 1). This is the direct consequence of the intense activity of researchers at the University of Neuchâtel during most of the last century. Their collections (as well as samples entrusted to them) were ultimately kept at the museum of natural history of this city, then transferred to the MHNG, which became a major repository for helminthological collections. Interestingly, only 22 species from this large pool are represented by types. These types are mostly from parasites of micromammals and have almost all been published either by Baer and collaborators in the 30s or by recent authors (e.g., Makarikov & Kontrimavichus, 2011). No parasites from birds were described in the country since Vasileva et al. (2000). Some material may have been registered without mention of their type status and it is possible, although unlikely, that other taxa from Swiss origin have been described without clear reference in foreign publications with their types conserved in collections not surveyed herein. Nevertheless, potential candidates for Swiss endemics should be looked for within those 22 species, especially amongst the micromammal parasites.
Host coverage
In comparison with Fuhrmann (1926) a significantly higher proportion of the Swiss fauna is currently known to host cestodes. The increase is particularly important for mammals (+130%) and to some extant for birds (+73%), while it is minor for other groups of vertebrates. Despite these figures, it should be noted that cestodes remain known from less than one third of the potential Swiss vertebrate hosts (see Table 2). Metacestodes have been found in 40 invertebrate taxa, an increase from 24 in Fuhrmann (1926), but still an extremely low number that is likely due to the paucity of recent life-cycles studies. Actinopterygii: There is a long tradition of fish parasitological studies in Switzerland and thus it is logical that the number of host species has only marginally increased since Fuhrmann (from 31 to 36). The tapeworm fauna from most common fishes is generally well known, although the problem of Proteocephalus spp. in whitefish remains unresolved (see above) and will necessitate detailed molecular studies to untangle. Nevertheless, despite the abundant literature and over a century of studies, the helminthes of the smaller or less common species remain poorly explored as tapeworms have been found in only 36% of the fish present in Switzerland (Table 2). An additional difficulty with fish is that introduced or invasive taxa are a problem in some waterbodies. A few of them have acclimated together with their parasite fauna, like the catfish Ameiurus melas (Rafinesque, 1820) and its proteocephalid Corallobothrium parafimbriatum Befus & Freeman, 1973.
Amphibia/Reptilia: Tapeworms are poorly diversified in herptiles, and the Swiss amphibian and reptile fauna is limited. Since Fuhrmann (1926) no new amphibian host has been found, but 2 snake and 1 lizard species have been discovered with cestodes. Tapeworms are probably present in a few more reptiles, but parasitological investigations of these hosts are particularly scarce. Aves: With cestodes described in a mere 20% of the 431 bird species recorded in Switzerland, this group of hosts is proportionally the most understudied, and consequently the largest reservoir of potential new species of parasites for the country's fauna. A large-scale study in many different countries worldwide showed that at least 40% of examined bird species hosted cestodes (Mariaux et al., 2017). In Switzerland, gaps are numerous as tapeworms have been found in only 35 out of 82 families of birds. Among the many families with no or very few recorded cestodes are small passerines [e.g., Acrocephalidae 0 species with cestodes out of 11 present, Hirundinidae (0/5), Phylloscopidae (1/11), Motacillidae (0/13) or Muscicapidae (2/23)]; however, some larger and very common birds, e.g., in Ardeidae (3/9) or Charadriidae (1/11) are also surprisingly understudied. It is for example highly unexpected that no tapeworm has been identified from the ubiquitous Ardea cinerea Linnaeus, 1758 in the country. Any basic parasitological survey of these often common and unthreatened birds would quickly add many species and dozens of host/parasites records to this checklist.
Mammalia: Most larger mammals, especially domestic ones, have been regularly studied and their parasitofauna can be considered as known. Starting in the 1960s regular surveys of micromammals have been undertaken and these hosts are now globally well covered too, resulting in cestodes described in 56% of the Swiss mammal fauna. Major gaps remain in Chiroptera with cestodes found in only 4 out of 26 species in the country.
Taxonomic problems
Confidence in cestode identifications reported herein may vary greatly among groups and depends obviously in part on whether recent revisions were made, or failing this, on the quality of initial determination. For many of the cyclophyllidean taxa, no recent taxonomic reassessment was performed, although a number of subgroups have been reviewed (at least in part) in the last two decades including several genera of Hymenolepididae (e.g., Vasileva et al., 1999, 2002; Makarikov & Georgiev, 2020) or of Dilepididae and Paruterinidae (e.g., Georgiev et al., 2004; Komisarovas et al., 2007; Dimitrova et al., 2017). Conversely, Bothriocephallidea and Oncoproteocephalidea have been more extensively reviewed e.g., by Scholz et al. (2007) or Kuchta et al. (2008). It remains that some of the older identifications may need confirmation. Part of the ancient material deposited in museums is, however, now over 100 years old and starting to deteriorate, making their study difficult. The development of molecular tools will certainly also bring new information on the composition and diversity of some taxonomically difficult groups, as recently exemplified by Brabec et al. (2023) but their use for identifying new taxa remain elusive for the time being as a comparative database is lacking.
Comparison with other European faunas
With 251 tapeworm species, the Swiss fauna is proportionally richer than that of other European countries, some of them much larger and with a marine fauna, like Finland (170 spp.), Iberian Peninsula (257 spp.), Slovakia (225 spp.), Poland (279 spp.) (Haukisalmi, 2015) or Italy (323 spp.) (Stoch, 2003). This observation stands even when considering only vertebrate hosts, as some of the above-mentioned reports did not consider invertebrates and metacestodes. For non-landlocked countries, the marine component of the specific diversity is obviously significant (e.g., Cyclophyllidea only count for 63% of the Italian specific diversity), but even lacking it, the Swiss fauna is remarkably diverse (61 spp./10'000 sq. km in Switzerland vs. 10.7 in Italy or 8.6 in Poland). By this metrics, Slovakia has the closest diversity (45.9) to Switzerland. Both countries share a landlocked situation at similar latitude, a mountainous landscape, as well as a strong helminthological tradition and therefore a higher number of species examined than in other countries. This last factor is likely the most significant, as also noted by Haukisalmi (2015).
More detailed comparisons of parasites faunas are hazardous as each country has a typical host diversity. However, Haukisalmi (2015, table 1, P6) provided a comparison of unique cestode species numbers per order of birds in various European countries to which we can compare figures for Switzerland. The latter are very similar to those in almost all countries considered, with the highest number of cestodes in Anseriformes, Passeriformes, Gruiformes, Galliformes and Podicepidiformes.
CONCLUSION
Despite lacking a marine fauna, Switzerland hosts a very rich cestode diversity that has been quite extensively studied, especially in the first half of the 20th century. Since the seminal synthesis of Fuhrmann (1926), our knowledge of tapeworms' diversity in the country expanded significantly, most notably in mammals.
This positive trend should, however, be put in perspective, as most of this progress was due to a very limited and quickly waning number of scientists. Presently, not only reports of new taxa are excessively rare and have all but stopped, but regular reports of known species in the wild fauna also became scarce. Among the factors leading to this situation, the vanishing training of taxonomists in this (and many other) group is certainly crucial. Furthermore, the taxonomy of animals like cestodes cannot rely on a population of amateur scientists to complement institutional research and the implementation of alternative (molecular?) systems of identification remain hypothetical at best given the poor comparative database available yet. This is worrying in a context of the threat to global biodiversity, and unfortunate because local taxonomic research would not require heavy investment. As demonstrated many times, including in Switzerland (e.g., Gigon & Beuret, 1991), a parasitological assessment of even the most common hosts would easily enrich the Swiss fauna and discover new indigenous taxa. This type of survey is, however, unlikely to flourish in the future as administrative agreements for collecting hosts, especially birds and some mammals, become increasingly arduous to obtain (Mariaux, 2021). As a result, today, and despite the number of prominent cestodologists who worked in Switzerland, less than one third of vertebrates in the country are known to harbor cestodes. The true extent of this parasitic fauna hence remains to be described.
Table 1.
List of cestodes and their hosts in Switzerland. Parasite species with a * and hosts with a ° were already reported in Fuhrmann (1926). L indicates that larval forms were found in this host. See “Material and Methods” section for details and abbreviations. Remarks are numbered in square brackets and are developed in the text.
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Table 2.
Hosts of cestodes in Switzerland
ACKNOWLEDGEMENTS
I am extremely thankful to Prof. Ian Beveridge (Melbourne), Dr Patrick Boujon (Lausanne), Prof. Boyko Georgiev (Sofia), Prof. Bruno Gottstein (Bern), Dr Roman Kuchta (Ceske Budejovice), Prof. Christian Lengeler (Basel), Prof. Tomas Scholtz (Ceske Budejovice), Prof. Gergana Vasileva (Sofia), and Dr Claude Vaucher (Geneva), who helped in various ways to improve this manuscript. I am grateful to Profs Janine Caira (Storrs, CT) and Kirsten Jensen (Lawrence, KS) for maintaining the Global Cestode Database at the University of Connecticut. Dr Voitto Haukisalmi (Helsinki) and an anonymous reviewer are thanked for their constructive suggestions.
I would also like to thank the following colleagues and curators who provided information on their collections: Dr Marco Bernasconi (Lucerne), Škoríková Blanka (Ceske Budejovice), Dr Danuta Cielecka and Dr Rusłan Sałamatin (Warsaw), Yvette Endriss and Dr Eduard Stoeckli (Basel), Dr Anne Freitag and Dr Michel Sartori (Lausanne), Dr Sophie Giriens (Fribourg), Eileen Harris (†) and Lauren Hughes (London), Dr Nicolas Kramar (Sion), Stephan Liersch (Chur), Dr Arseny Makarikov (Novosibirsk), Dr Nicolas Margraf (La Chaux-deFonds), Dr Lucia Pollini (Lugano), Dr Martina Schenkel (Zurich), Dr Sabrina Schnurrenberger (Winterthur), Dr Denis Vallan (Aarau).
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited (see https://creativecommons.org/licenses/by/4.0/).
BIBLIOGRAPHY
Appendices
Annex 1: List of specimens in collections. Catalogue numbers without collection reference are from the Muséum d'histoire naturelle de Genève (MHNG-PLAT-). Type status is indicated with HOLO (Holotype), LECTO (Lectotype), PARA (Paratype), SYNT (Syntype) or TYPE (Type of unknown status).
BOTHRIOCEPHALOIDEA, TRIAENOPHORIDAE, Bathybothrium rectangulum 27276, 40291, 55791, 55798, Eubothrium crassum 17858-9, 19002, 19327, 23873, 28077-8, 28080-1, 38317, 40808-9, 88297-302, Eubothrium salvelini IPCAS H02/1, IPCAS C126/12, 27278, 29413, 38313, 38364, 33625, 36722-6, 36728-9, 55807, 82342-4, 82660, ZMZ-122912, Triaenophorus crassus 42479-81, 55808, 57528, 57532, NHM 1928.1.9.130-134, Triaenophorus nodulosus 11607, 18170, 18498, 27937, 36003-7, 38257, 38262, 42482-6, 54161, 54426, 57662, 57667-8, 63396-8
CARYOPHYLLIDEA, CARYOPHYLLIDAE, Caryophyllaeus fimbriceps 78801-3, LYTOCESTIDAE, Caryophyllaeus laticeps 18338, 27277,38331, 39225, 40289, 70965, 71127, 78800, 78837, 78840-4, USNM 1355422-3, Caryophyllaeides fennica 78797, NHM 1928.1.9.202-203, USNM 1355424-5, Khawia baltica 78804-5
CYCLOPHYLLIDEA, AMABILIIDAE, Joyeuxilepis acanthoryhncha 42356, ANOPLOCEPHALIDAE, Anoplocephala magna 38376, 56118, Anoplocephala perfoliata 40241, 56079, Anoplocephaloides dentata 17608, 18408, 30635, 41787, 82353, 82370, 82372, 82396-7, Atriotaenia incisa 14620, 57153, 57180, Ctenotaenia marmotae 130473, 27280, 38616 (HOLO of Cittotaenia avicola), 38617, 37276, 38332, 40497-8, 40500-1, Equinia mamillana 41792-4, 56113, Eurotaenia gracilis 11430-2 PARA (of Paranoplocephala gracilis), 11583, 19182, 38187, 82345, 82349, 82351, 82374-5, Genovia wimerosa 41803-4, Microticola blanchardi 13482, 82346, Moniezia benedeni 38304, 41600-1, 56047, 56059, Moniezia expansa 41588-9, 57237, Mosgovoyia pectinata 40503, 40508, 57115, 57171, Neoctenotaenia ctenoides 40489, 56164, 56167, Oochoristica rotundata 41696, Paranoplocephala omphalodes 12153, 12166, 12217, 13857, 17742, 17771, 20000, 38186, 40078-9, 41795-6, 41799-800, 82378, 82383, Thysaniezia giardi 40898-900, Thysanosoma actinioides 42471, CATENOTAENIIDAE, Catenotaenia dendritica 40369-70, Catenotaenia henttoneni 17637-8, 18361, 18368, 39305 39378 39446, Catenotaenia pusilla 37655, 40379-81, Skrjabinotaenia lobata 12162, 17625-8, DAVAINEIDAE, Davainea andrei 40620 SYNT, Davainea proglottina 27994, 28053, Davainea tetraoensis 55227, Fernandezia spinosissima 18326, 32733-5, 77626, Idiogenes flagellum 27997, Paroniella urogalli 27997, Raillietina frontina 42078, Raillietina tetragona 27315, 27984, Skrjabinia bonini 42034, 55973, NHM 1928.1.6.107-116, USNM 1318063, USNM 1348473 SYNT of R. columbae, Skrjabinia cesticillus 27986, 28076, DILEPIDIDAE, Anomotaenia brevis 13476, Anomotaenia cyathiformis 40119, Anomotaenia dehiscens 27910, 40121-2, NHM 1928.1.9.43-48, Anomotaenia microphallos 40154, 40157, 56925, Anomotaenia microrhyncha 39308, Anomotaenia stentorea 39307, Burhinotaenia coronata 41822, Choanotaenia orioli 40455 SYNT, 40459, Choanotaenia passerina 15350, 39309, Dictymetra sp. 50022, Dilepis cypselina 40638, Dilepis undula 11435, 11494, 11608, 12161, 13400, 13475, 15348, 17736, 17614, 17824, 18434, 18553, 27940, 27970, 32725-32, 32767-72, 38279, 38888, 38904, 38942, 38963, 39386, 39394, 40666-7, 40670, 77627-30, hepatocestus hepaticus 11483, Liga sp. 50023, Molluscotaenia crassiscolex 11380, 11394, 11399, 11413, 11420, 11480-1, 11485, 11489-90, 11578, 11581, 11613, 14289, 17743, 17749-51, 17765, 18174, 18178, 18219, 18229, 18232, 18234, 18245, 18378, 18429, 18478, 18481, 18483, 18485, 18552, 30630, 30640, 30661, 30687, 30808, 30899, 38884, 38887, 38892, 38896, 38899, 38903, 38922, 38930, 38932, 38935, 38941, 38943, 38948, 38952, 38962, 38967, 38970, 38972, 38978, 38981, 38988, 38992, 38996, 39001, 39005, 39011, 39017, 39026, 39031-2, 39322, 39331, 39365-6, 39368, 39373, 39376, 39379, 39384-5, 39390, 39392-3, 39397, 39400, 39402-3, 39406, 39408, 39410, 39417, 39419-20, 39438, 39440, 40440, 41623, 48316, 82357, 82379, Monopylidium albani 32776, Monopylidium crateriformis 27295, 40443-4, 40441-2, Monopylidium galbulae 40130, Monopylidium musculosa 32775, 40452, Multitesticulata filamentosa 13776, 17823, 18367, 39303, 39480, 41613-4, Neoliga depressa 27908-9, 39313, 40123-4, Paricterotaenia porosa 18377, 27929, 27945, 40821, 41854, 41856, NHM 1928.1.9.170-175, Platyscolex ciliata MUW-114097, Pseudoangularia sp. 40123, Sobolevitaenia spinosocapite 32724, 32773-4, 38278, 57134, Sobolevitaenia verulamii 13401, 40117, Spassspasskya passerum 11434, 13474, 40178, Spiniglans affinis 32745, Spiniglans constricta 27964, 28086, Emberizotaenia raymondi 32503 HOLO, 32504 PARA, 82652 PARA (of Unciunia raymondi), DIOICOCESTIDAE, Dioicocestus asper 40680, DIPYLIDIIDAE, Dipylidium caninum 17864, 18343, 28082, 28084, 40747-8, 40751, 40753, 40755, 56123, GRYPORHYNCHIDAE, Paradilepis scolecina 41776-8, HYMENOLEPIDIDAE, Anatinella kazachstanica 57479-80, MUW-114104, MUW-114107, Aploparaksis cirrosa 27930, Aploparaksis crassirostris 27941, Aploparaksis filum 27312, 38294, 40878-9, Aploparaksis furcigera 40887-8, 56550, Armadolepis (A.) jeanbaeri 17611, 39226-9 PARA, 41189 HOLO, 41190 PARA, Armadolepis (B.) myoxi 13858-5, Arostrilepis horrida 13867-8, 18514, 19293, 19309, 19664, 30491, 30553, 41097, Arostrilepis janickii 13866 PARA, 18499 PARA, 18513 PARA, 19291 PARA, 19300 PARA, 19308 PARA, 19662 PARA, 41096 HOLO, Cladogynia guberiana 15589, 57464, 57467, MUW-70736, MUW-70750-6, Cladogynia macracanthos 41125-9, 55608 SYNT (of hymenolepis macracanthos), NHM 1928.1.9.33-42, Cladogynia serrata 41270, Cloacotaenia megalops 41144, MUW-114116, Confluaria furcifera 49190, Confluaria multistriata 27934, NHM 1928.1.9.66-73, Confluaria pseudofurcifera 18328, 40964, 40971 PARA, 41076 PARA, 41077 HOLO, 41236 PARA, 41289, 124885, Coronacanthus integrus 15351, 17755-6, 30675, 39029-30, 39033-4, 39041, 39043, 39289, 39292, 41090, 41100-2, 41210, Coronacanthus omissus 39027, 39288, 39290-1, 41210-11 SYNT, 49031, Coronacanthus integrus 41237 SYNT (of hymenolepis polyacantha), Cryptocotylepis globosoides 10736, 19290, 39317, 41084, Dicranotaenia coronula 27975, 28089, 41001-3, Diorchis acuminata 40683, 40694, 55782, MUW-114109, Diorchis ransomi MUW-71581-8, MUW-114111, Diploposthe laevis 55640, Ditestolepis diaphana 11389, 11408, 11415, 11418, 11476, 11488, 11587, 17778, 18231, 18233, 18382, 18535-8, 18851, 26382, 26384, 30663, 30666, 30678, 30805-6, 36339, 38881, 38894, 38907, 38913, 38920-1, 38926, 38940, 38945-6, 38955, 38966, 38976-7, 38985, 38990, 38995, 38999, 39009, 39015-6, 39020, 39022-3, 39267, 39320, 39362, 39381, 39388, 39399, 39412, 39414, 39435, 39694, 41015, 82356, Dollfusilepis hoploporus 40964, Dubininolepis rostellata 41251-4, 41256, 56554, Echinocotyle anatina 27983, NMB-CEST-00039a, Echinolepis carioca 40974, Fimbriaria fasciolaris 39310-11, 40820, 55624, MUW-50053, Gulyaevilepis tripartita 18226, 18382, 26382, 38891, 38921, 38977, 39009, 39016, 39022, hispaniolepis villosa 27303, 55742 NMB-CEST-00033a, hymenolepis armata 55765, hymenolepis capillaroides 40968, hymenolepis diminuta 17735, 30801, 41018, 41021, hymenolepis hibernia 27286, 27927, 27938, 41168, hymenolepis murissylvatici 17633-6, 18524-6, 39279, 39281, 39284, 39286, 41187-8, hymenolepis simulans 44405, hymenolepis sp. 41228, hymenolepis sulcata 13723-32, 13777 13854, hymenolepis teresoides 41319, hymenolepis uliginosa 27926, hymenolepis uncinata 17808-11, 39364, Lineolepis scutigera 11378, 11382, 11386, 11410, 11412, 11416, 11475, 11492, 11501, 11579-80, 11600, 17761-3, 18546, 30667, 38890, 38895, 38898, 38914, 38917, 38925, 38929, 38934, 38938, 38956, 38997, 39002, 39007, 39013, 39021, 39264, 39268, 39326, 39330, 39353, 39374, 39377, 39380, 39396, 39411, 39431, 39441, 38910, 41326 SYNT (of hymenolepis toxometra), Microsomacanthus abortiva 40916, 40918, 55634, Microsomacanthus arcuata 40933, 41344, 55621, 55771, Microsomacanthus collaris 27943, 40984-6, Microsomacanthus compressa 40999, Microsomacanthus microcephalus 27953, Microsomacanthus microsoma 37989, Microsomacanthus paracompressa MUW-114092-4, MUW-114101, MUW-114112-3, MUW-114117-9, MUW-114123, Microsomacanthus setigera 41273, 41278, Microsomacanthus spiralibursata MUW-114090-1, MUW-114096, MUW-114099, MUW-114114, MUW-114120, MUW-114122, Milina grisea 14781, 18534, Monorcholepis dujardini 11585, Neomylepis magnirostellata 18390, 30676, 39036, 41134 SYNT, Neoskrjabinolepis merkushevae 11417, 11421, 11504-5, 11601, 11779, 17974, 18227, 18235, 38905, 38911, 39369, 39418, Neoskrjabinolepis schaldybini 11383, 11387, 11391, 11396-7, 11403, 11482, 11484, 11493, 11497-8, 11571, 11576, 11590, 11592, 11596-7, 11599, 11602, 11605-6, 17746, 17760, 18173, 18176, 18230, 18380, 18431, 18433, 18480, 18506, 18547-8, 30602, 30604, 30639, 30662, 30665, 30802, 30807, 36338, 36732, 38882, 38897, 38902, 38908-9, 38919, 38924, 38928, 38937, 38944, 38954, 38958, 38960, 38969, 38975, 38980, 38984, 39025, 39266, 39316, 39327, 39329, 39349, 39354, 39367, 39383, 39401, 39437, 39442, 41261-2, Neoskrjabinolepis singularis 11379, 11385, 11392, 11395, 11401, 11404, 11406, 11499, 11574, 11577, 11588, 11594, 11598, 11610, 39018, 39351, 39356, 39426, 39439, Parabissacanthes bisacculina 40952, Parabissacanthes philactes 15588, 57471, 57478, MUW-114103, MUW-114105-6, MUW-114121, Passerilepis brevis 32750-1, Passerilepis crenata 11433, 12160, 13399, 27965, 27978, 32736-7, 32742-3, 32779-83, 32843, 35742-4, 41013, 41264, Passerilepis passeris 15603, 32748-9, 32784-6, 41069-70, Passerilepis stylosa 27291, 27985, 41313, 41315, 41317, 55927, Pseudhymenolepis redonica 15471, 17754, 17819, 18387, 18554-5, 42014 SYNT, Rodentolepis asymmetrica 11779-81, 11785-6, 17613, 18156, 18165, 18362-4, 18365, 18399, 18502-3, 18509, 18511, 19075, 19078, 19162-4, 19181, 19183-5, 19203, 19205-6, 19321, 19328, 19663, 30533, 30549, 30668-9, 30672-3, 39304, 39413, 40938, 40940, 41308, 82387, 129733, Rodentolepis erinacei 41032, Rodentolepis fraterna 41063-7, Rodentolepis microstoma 15346, 37496-503, 37507, 37509, 37511, 37644-51, 41164, 41167, 41172, Rodentolepis straminea 11565, 17612, 18508, 18517-22, 18532, 35297, 39294-5, 82473, Skrjabinacanthus jacutensis 11496, Sobolevicanthus fragilis 41059, Sobolevicanthus gracilis 27911, 27981-2, 41087, MUW-70552-4, MUW-114100, MUW-114102, Sobolevicanthus gracilissimus MUW-70631, Sobolevicanthus krabbella MUW-114095, Soricinia globosa 19292, 41083 SYNT (of hymenolepis globosa), Soricinia infirma 11384, 11414, 11419, 11503, 11591, 24135, 26360, 30804, 38947, 38961, 38987, 38991, 39000, 39010, 39328, 39361, 39371, 39415, 39429, 39433, 39436, 82355, Staphylocystis alpestris 40924 SYNT (of hymenolepis alpestris), Staphylocystis brusatae 17816-8, 39407, Staphylocystis furcata 11400, 11423, 11486, 11495, 11500, 11573, 11595, 11612, 18175, 18217, 18258, 18430, 18504, 18510, 18540, 30555, 38900, 38973, 39398, 39409, 39427, 41073-5, Staphylocystis pistillum 17739, 17753, 17759, 17785, 17787-8, 17790, 18542-3, 29656, 36340, 41075, 41231-3, 41235, 42506, 49191, 49248, 49250, Staphylocystis scalaris 17740, 17780-4, 17786, 18253, 41260, Staphylocystis tiara 17741, 17801-7, 18435, 18544-5, 39280, 39375, 39421, 41322, Staphylocystoides stefanskii 11388, 11390, 11398, 11407, 11422, 11481, 11572, 11593, 18479, 18539, 18549, 38906, 38912, 39004, 39265, 39430, 41303-4, Triodontolepis bifurca 18257, 39028, 39283, 39293, 40945, Triodontolepis hamanni 10737, 17748, 39282, 41089, Urocystis prolifer 11479, 11487, 11502, 11604, 17745, 17764, 17777, 18177, 18220, 18259, 18379, 18550-1, 18553 18842, 30637, 30664, 30674, 30677, 30803, 38893, 38918, 38939, 38951, 38965, 38986, 38989, 38994, 38998, 39003, 39014, 39019, 39318-9, 39325, 39350, 39370, 39382, 39389, 39395, 39416, 39432, 39434, 39443, 41196 SYNT (of Vampirolepis neomidis), 48042, 48315, Vampirolepis baeri 18329 HOLO, 27230, Vampirolepis balsaci 14780, 14782, 39341-3, 39345, 40943, Vigisolepis spinulosa 11381, 11393, 11402, 11405, 11409, 11411, 11575, 11589, 11603, 11611, 18218, 18228, 18381, 18432, 30554, 38883, 38885, 38889, 38901, 38916, 38923, 38927, 38931, 38933, 38936, 38950, 38953, 38957, 38959, 38968, 38974, 38979, 38983, 39006, 39012, 39024, 39315, 39321, 39323-4, 39352, 39391, 39404, 41301-2, Wardium arciminosa 18533, 27282, 27285, 27974, 39306, Wardium recurvirostrae 27287, 27939, 27972, Wardoides nyrocae cygni MUW-114108, MUW-114110, MESOCESTOIDIDAE, Mesocestoides lineatus 14550, 14562, 14580, 14582-94, 14601-2, 14604-5, 14607-11, 39339, 41539, 41541, 45076, Mesocestoides litteratus 27912-3, Mesocestoides perlatus 27310-1, Mesocestoides sp. USNM1397704, Nematotaenia dispar 27931, 38025, 38285, 40599, 41638, PARUTERINIDAE, Anonchotaenia globata 32747, 40218, 41869, Cladotaenia cylindracea 39312, 40524, 40528, Cladotaenia globifera 27936, 27948, 32744, Neyraia intricata 41647, Notopentorchis sp. 137308, Orthoskrjabinia bobica 18327, 32778, Orthoskrjabinia conica 32746, 32777, Spasskyterina trianguloides 17596, TAENIIDAE, Echinococcus granulosus 38321, 40789-90, 43510-1, Echinococcus multilocularis 14544, 14581, 40789-90, 42312, 42316, 43510-1, hydatigera taeniaeformis 13765-75, 13856, 14230, 14276, 14628, 17606-7, 17772-3, 18210, 18512, 27254, 31071, 32752, 37387, 37389-91, 37401, 38216, 39285, 39287, 42350, 48040,, Taenia crassiceps 14272, 14279, 14534, 14536, 14538-42, 14546-9, 14551, 14554-6, 14558, 14561, 14563, 14565-71, 14574-5, 14577-9, 14599, 17605, 18180, 18336, 18351, 24478, 27900, 27942, 27966, 37388, 38214-5, 38219, 38222, 38318, 42255-61, Taenia martis 14637-66, 14716-9, 14726, 15355, 15414-6, 15428-9, 17601-2, 17737-8, 17868-70, 18507, 18528-9, 24334, 30580, 41234, 42290-2, 42294, Taenia multiceps ZMZ-G224, Taenia pisiformis 14552-3, 38263, 38265, 40605, 42312-3, 42315-6, Taenia polyacantha 14535, 14537, 14543, 14557, 14564, 14572-3, 14576, 15545, 14600, 17603, 17861-2, 28162, 28166-7, 30506-7, 39230-1, 41237 SYNT, 42317, 42319, 48333, 56818, Taenia saginata MHNF-5849, 42328-9, 38268, 38270-2, 57166, NWSW-13092, ZMZ-122915, ZMZ-122796, ZMZ-122761, Taenia serialis 41016, Taenia solium 42252, 42338, 57273, ZMZ-120573, Versteria mustelae 17600, 24333, 28085, 38337, 42300-2, 423614, 58867
DIPHYLLOBOTHRIIDEA, DIPHYLLOBOTHRIIDAE, Diphyllobothrium latum 18383, 18385, 27273, 27995, 38371-2, 40708-11, 43073-5, 43079-80, 56260, 57124, 82431, USNM 1348495-6, Ligula digramma 38380, Ligula intestinalis 17865, 38302, 38305, 38309, 38312, 38315, 41508, GBIFCH00596771, NMB-CEST-00009b, ZMZ-120412, ZMZ-120577, Schistocephalus solidus GBIFCH00596744
ONCOPROTEOCEPHALIDEA, PROTEOCEPHALIDAE, Corallobothrium parafimbriatum 32994, Glanitaenia osculata IPCAS C49/1, 67699-700, 68395-7, 84707-10, 84712, 91260, 91839-42, Ophiotaenia europaea 49149, Proteocephalus filicollis 27302, 41985, Proteocephalus longicollis 15601, 16920-3, 16925, 19239, 19278, 19280-4, 19667-8, 21681, 27301, 38353, 41529, 41353-4, 41356, 41358, 41367 SYNT (P. salmonisumblae), 86982, NHM 1998.2.178.42, NHM 1998.2.20.1-2, NHM 1998.5.14.4, USNM 1348661 TYPE, USNM 1348668 LECTO, USNM 1382490, USNM 1382832, USNM 1382933, USNM 1395121, Proteocephalus percae IPCAS C29/1, 16924, 19238-45, 19268-77, 19279, 19285-8, 27300, 28749-50, 36744-5, 39479, 41357, 41363-5, 54160, 57357, 61489, 63221, 63395, 63399-400, NHM 2000.1.25.19, NHM 2000.6.1.1-2, USNM 1348656 (TYPE of P. dubius), USNM 1382102, USNM 1382827, USNM 1385360-1, Proteocephalus torulosus IPCAS C32/1, 19666, 27916, 41368
SPATHEBOTHRIIDEA, ACROBOTHRIIDAE, Cyathocephalus truncatus 15600, 32823, 38330, 40579-80, 55813, 70959, 70960-1, 70963, 88303
TETRABOTHRIIDEA, TETRABOTHRIIDAE, Tetrabothrius (T.) macrocephalus 42432, 55979
