An annotated catalogue of the tongue worms (Pentastomida) held in the Museum fçr Naturkunde Berlin

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1 Zoosyst. Evol. 86 (1) 2010, / DOI /zoos An annotated catalogue of the tongue worms (Pentastomida) held in the Museum fçr Naturkunde Berlin Doreen Ræhlig 1, Jason A. Dunlop*, 1, Jos Horacio Grau 1, 2 and Anja Friederichs 1 1 Museum fçr Naturkunde, Invalidenstraße 43, Berlin, Germany 2 Departamento de Zoologia, Universidade de S o Paulo (USP), Rua do Mat o, trav. 14, no 321, Cidade Universit ria, S o Paulo, SP, , Brazil Abstract Received 15 September 2009 Accepted 28 October 2009 Published 17 March 2010 Key Words ZMB nomenclature taxonomy phylogeny parasite Heymons An annotated catalogue of the type and non-type material belonging to the enigmatic arthropod group Pentastomida (tongue worms) in the Museum fçr Naturkunde Berlin is presented. Aspects of human infection, current ideas on their affinities and some recent developments in the study of these animals are briefly reviewed. Types of eight species have been recorded: Raillietiella affinis Bovien, 1927, Raillietiella bicaudata Heymons & Vitzthum, 1935, Pentastomum javanicum Bovien, 1927, Raillietella kochi Heymons, 1926, Elenia lialisi Heymons, 1939a, Raillietiella mabuiae Heymons, 1922, Raillietella shipleyi Heymons, 1926 and Cephalobaena tetrapoda Heymons, Five of these types are currently considered valid species. According to the literature, type material from Elenia travassosi (Heymons, 1932) should be in Berlin, but could not be traced during the present review and may have to be considered lost. A further twentysix non-type species were recorded, plus specimens assigned to two names currently regarded as nomina dubia. This yields a total of 31 currently valid species in Berlin, representing nearly 25 % of the known world fauna. Much of the Berlin material derives from the important Richard Heymons collection, but significant historical specimens collected in Brazil in the early nineteenth century by Johann Natterer was also discovered during the course of this project. Introduction Tongue worms (Pentastomida) are an unusual group of parasitic arthropods. The ca. 130 living species (Almeida & Christoffersen 1999) are found as adults exclusively in the respiratory tracts of vertebrates. They typically occur in reptiles, but have also been recorded either as adults or larvae from amphibians, birds and mammals. Some of the range of both definitive and intermediate hosts is detailed in the collection data presented in our catalogue. They occasionally infect humans (Haffner 1974; Fain 1975; see also below), although man is not their natural host. Generally, the economic impact of tongue worms is fairly low and this is reflected in a relative paucity of studies devoted to this largely tropical group. Self (1969) published a comprehensive bibliography of studies of these animals up to that date. Rudolphi s (1819) name Pentastomida (five openings) derives from an early misinterpretation of the two pairs of hooks flanking the mouth as additional mouth openings, while their common name tongue worm reflects the distinctly tonguelike shape of some species, such as those in the genus Linguatula Frælich, The adult body, which is typically a few centimeters long but can reach up to 14 cm, is highly annulated and in other taxa is more worm-like and can have a distinct head region (e.g. Kiricephalus Sambon, 1922). Many organ systems are lacking and summaries of pentastomid biology including their life cycle, definitive and intermediate hosts, and adaptations to parasitism can be found in, e.g., Heymons (1935), Riley (1986) or Haugerud (1989), and references therein. The aim of this present paper is to summarise both the type and non-type pentastomid material held in the Berlin museum as well as to briefly review their role as human parasites, together with recent developments towards resolving their likely phylogenetic affinities. * Corresponding author, jason.dunlop@mfn-berlin.de

2 130 Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Richard Heymons An important historical figure in the study of pentastomids was the Berlin zoologist Richard Heymons ( ). Born in Berlin, where he also studied, Heymons (Fig. 1) was invariably associated with the Berlin University; except for a brief period ( ) in Hannoverisch-Mçnden. In 1906 he received his professorship in Berlin and at the same time worked as one of the entomology curators at the university s zoological museum (now the Museum fçr Naturkunde). In 1915 he was appointed to a chair at the agricultural veterinary faculty, which he held until his retirement in Further biographical information can be found in Ulrich (1961). Over the years Heymons made a number of key contributions to the biology and systematics of tongue worms (Heymons 1922, 1926a b, 1928, 1930, 1932a c, 1933, 1935, 1939a b, 1940a c, 1941a c, 1942, 1943; Heymons & Vitzthum 1935, 1936), including the description of a number of new species (Tab. 1). Although Heymons no longer worked in the Berlin museum at the time he wrote his pentastomid papers, a survey of the Berlin museum s arachnid and myriapod collection revealed a number of his types one from a paper published together with Hermann Graf Vitzthum ( ) together with further material derived from his studies of the group which is clearly indicated by Coll. Heymons. on the labels. Seventy-five lots from the Heymons collection should be present, and all but five (numbers 14, 15, 26, 67, 75) Figure 1. Richard Heymons ( ), who gathered together much of the MfN s pentastomid material. Image courtesy of the Museum fçr Naturkunde Berlin, Historische Bildund Schriftgutsammlungen (Sigel: MfN, HBSB). Bestand: Zool. Mus. Signatur: B I/69 (Portrait of R. Heymons). could be matched to material currently in the Berlin museum. Note that other Heymons types were deposited in the Hamburg Museum and the London School of Hygiene and Tropical Medicine (Tab. 1). The collection data reveals that Heymons received much material from the Berlin Zoo and Aquarium, at Table 1. The twelve pentastomid species/subspecies described by Richard Heymons and their type repositories according either to the present study or the literature. Taxa arranged in their approximate phylogenetic sequence. Taxon Repository Remarks Cephalobaenidae Cephalobaena tetrapoda Heymons, 1922 Berlin this study Raillietiellidae Raillietiella bicaudata Heymons & Vitzthum, 1935 Berlin? see text Raillietiella boulengeri spiralis Heymons, 1939b* London** Raillietiella kochi Heymons, 1926a Berlin/Hamburg this study; Weidner (1959) Raillietiella mabuiae Heymons, 1922 Berlin this study Raillietiella shipleyi Heymons, 1926a Berlin this study Sebekidae Alofia travassoi Heymons, 1932b Berlin lost? Elenia australis Heymons, 1932a Hamburg cf. Weidner (1959) Elenia lialisi Heymons, 1939 Berlin this study Porocephalidae Armillifer armillatus intermedius Heymons, 1940b London** Armillifer moniliformis australis Heymons, 1932*** Hamburg cf. Weidner, (1959) Waddycephalus vitiensis Heymons, 1932a Hamburg cf. Weidner (1959) * The publication date is often cited as 1940 the overall year of volume 11 of Zeitschrift fçr Parasitenkunde but the first issue which also contains Heymons paper bears the inscription Abgeschlossen 13. Juli 1939 ; later issues came out in ** Cited as the London School of Hygiene and Tropical Medicine (Prof. P. A. Buxton) we cannot guarantee that this is where the material currently resides, as there are citations to Buxton in Liverpool too. *** This subspecies name appears to us to be problematic. Although Heymons (1935) refers to his own name as dating from 1932, we could not trace it in any of the three 1932 papers (see also Self 1969) we are aware of. museum-zoosyst.evol.wiley-vch.de

3 Zoosyst. Evol. 86 (1) 2010, least partly through Professor Max Koch, as well as material from the Hamburg Zoo. Heymons was also involved in exchanges with the Natural History Museum in London, the Instituto Butantan in S o Paulo and the Zoological Museum (Natural History Museum of Denmark) in Copenhagen. Prosper Louis Bovien (1894?) was based at an agricultural faculty in Copenhagen from the late 1920 s and could conceivably have been the source of the latter material, as well as two further syntypes from Bovien (1927) which were identified in the Berlin collection during the present study. Heymons also appears to have obtained material from Vienna derived from collections by Natterer : presumably the naturalist and explorer Johann Natterer ( ) who took part in a major Austrian collecting trip to Brazil (summarised by Riedel-Dorn 2000). Natterer s Brazilian specimens were extensively used by the Vienna-based helminthologist Karl Moriz Diesing ( ) in some of his key early studies of pentastomids (e.g. Diesing 1835). It should be noted that Heymons & Vitzthum (1936) make extensive reference to having studied the Natterer collection in Vienna, but there is no indication on the Heymons/Berlin labels whether these speciens represent loans, gifts or exchanges, or whether they should be even be considered part of Diesing s syntype series for the species in question. Nevertheless, the discovery of what must at least be regarded as voucher material from this early period of pentastomid research is an important discovery for the Berlin collection. Human infection While human infection by pentastomids is typically cited as rare, and largely restricted to warmer regions today, historically they seem to have been rather more widespread even in Europe. Koch (1906) reported a 12 % rate of visceral linguatulosis derived from around 400 adult autopsies in Berlin around the turn of the century. Sonobe (1927), again in Berlin but a few years later, recorded a 3 % infection rate based on autopsies of the liver. While subsequently becoming rarer in Europe (e.g. Symmers & Valteris 1950; Drury 1962), there is a substantial modern literature about infection in the Middle East, Africa and Asia in particular. Parasitism by pentastomids may be locally more common than is widely appreciated. Infection rates, based on adult human autopsies, of 8 % in Cameroon and 23 % in Zaire have been cited (Mapp et al. 1976, and references therein). A remarkable 45 % rate was documented for Malayan aborigines (Prathap et al. 1969), while Yagi et al. (1996) reported that up to 20 % of the human population in some parts of Sudan could be affected by the pentastomid-mediated Marrara syndrome (see below) at some stage of their lives. Dermatitis induced by a juvenile Sebekia sp. has been reported (Mairena et al. 1989), but Linguatula and Armillifer are the genera normally associated with human infection (Fain 1975). The usual clinical terms are linguatulosis, pentastomosis or porocephalosis; largely depending on the taxon responsible. Humans are typically accidental hosts and, given that they are rarely consumed by reptiles or jackals, they represent a dead end intermediate host who does not transfer the infection further. Riley (1986) provided a detailed review of human interactions with pentastomids and valuable modern accounts of infection pathology, case histories and summaries of the relevant literature can be found in, e.g., Tappe et al. (2006), Yao et al. (2008) and Tappe & Bçttner (2009). The way in which pentastomids in general evade the body s immune response were reviewed by Riley (1992). While most patients are asymptomatic, coughing, night sweats and abdominal pain have been reported; symptoms depending on whether infection is by adults or nymphs and thus where in the body the pentastomids have become established. Nymphs usually die in the host tissue after a couple of years; after which they can sometimes calcify, yielding a characteristic crescent shape when X- rayed (Bretland 1962; Mapp et al. 1976). A heavy parasite burden can have serious clinical consequences (Pan et al. 2005) and in a very few cases ends fatally (Yapo Ette et al. 2003; and references therein). Linguatulosis is often associated with contact to dogs dogs, jackals and other carnivores are the usual final host which might explain its historical prevalence in temperate urban environments. In the Middle East and South East Asia infection rates in stray dogs of between c % have been reported (Meshgi & Asgarian 2003; and references therein). Two forms of infection can occur (cf. Schacher et al. 1969). Visceral linguatulosis would normally affect a ruminant intermediate host (e.g. Shekarforoush et al. 2004; Ravindran et al. 2008; and references therein) and involves picking up eggs sneezed, or defecated, out by the definitive host. Once in the intestines of the intermediate, the eggs hatch into larvae, which migrate into other internal organs (particularly the liver) where they form cysts or granulomas. This can also occur in humans, even today (Machado et al. 2006), even though the host may not be aware of the infection. Alternatively, in nasopharyngeal linguatulosis juveniles already encysted in an intermediate host (usually a domestic herbivore) are consumed. Raw or poorlycooked meat, particularly liver or lymph nodes, seems to be the definitive route of transmission to humans. Within minutes to hours, the nymphs are liberated from their cysts in the stomach and migrate up the oesophagus and enter the airways (nose, pharynx or lungs), where they attach themselves with the hooks flanking the mouth (Kahil & Schacher 1965; Siavashi et al. 2002). This can result in patients suffering headaches, coughing, nasal discharge, etc. Sneezing out eggs obviously mediates the further spread of infection. This hypersensitive reaction to the parasite may be mediated by proteins (Buslau et al. 1990) and can be treated with antihistamines. Nasopharyngeal linguatulosis is often museum-zoosyst.evol.wiley-vch.de

4 132 referred to as Halzoun syndrome in the Middle East, since it typically manifests itself after religious festivals during which raw meat is traditionally consumed. In Sudan it is known as Marrara syndrome, whereby Marrara is a popular local dish prepared from raw offal (Yagi et al. 1996). Larvae of both Lingulatula and Armillifer have occasionally been reported as having presumably accidentally migrated into the human eye; see e.g. McKie Ried & Ellis Jones (1963), Lang et al. (1987), Lazo et al. (1999) and references therein. By contrast, infection with Armillifer is usually associated with contact to snakes, and humans are only ever intermediate (or visceral) hosts affected by encysted larvae. Eggs can be ingested via raw or poorly-cooked snake meat (Mçlder 1989; du Pleiss et al. 2007; Yao et al. 2008) and infection is thus more prevalent in those African and Asian countries where such practices are common. Alternatively, Dakubo et al. (2008) described how in West Africa infection could be related to clans who use the python as a totem; close contact increasing the risk of picking up eggs from the definitive host via slime or faeces. Affinities Of particular interest is the long-standing debate about the phylogenetic affinities of these enigmatic animals (Self 1969; Riley et al. 1978). The numerous, possibly reductive, characters associated with pentastomids parasitic lifestyle only serve to complicate the issue. Historically, tongue worms were initially compared to various groups of parasitic worms (Frælich 1789; Rudolphi 1819). Once the arthropod-like nature of their cuticle was recognised, Schubart (1853) and Leuckart (1860) drew parallels between pentastomid larvae and Acari (mites and ticks). This concept was strongly influenced by their superficial resemblances to gall mites (Eriophyidae) which also have a rather elongate, annulate body and only two pairs of legs. Tongue worms were also compared historically to myriapods and/or to the stem-arthropod groups Onychophora (velvet worms) and Tardigrada (water bears) (von Haffner 1977). Although, the arthropod nature of their larvae was demonstrated by Osche (1959), they were still been interpreted by some authors as a wholly independent group (e.g. Self 1969), either sharing a common ancestry with annelids and arthropods or as somehow being intermediate between these two major animal groups, perhaps with closer affinities to worms (e.g. Heymons 1935). The pentastomid fossil record extends back some 500 million years to the late Cambrian (Walossek & Mçller 1994; Walossek et al. 1994) and the current literature offers essentially two competing phylogenetic hypotheses; see e.g. Lavrov et al. (2004), Waloszek et al. (2006) and Almeida et al. (2008) for recent reviews, albeit coming to different conclusions. Lavrov and collagues argued that pentastomids are derived crustaceans, probably close to Branchiura (fish lice). The crustacean hypothesis has Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) its origins with van Beneden (1849), who proposed affinities between the larvae of pentastomids and the parasitic copepod genus Clavella [as Anchorella]. In its modern form it is based largely on sperm morphology and developmental characters (Wingstrand 1972; Riley et al. 1978), and branchiuran affinities have been strongly supported by a series of molecular studies (Abele et al. 1989; Petersen & Eernisse 2001; Lavrov et al. 2004; Møller et al. 2008). In detail, Lavrov et al. sequenced the entire mitochondrial genome and their conclusions were thus supported both by mtdna and the rearrangements within the mt genome, while Møller et al. s broader taxon sampling allowed them to demonstrate that pentastomids resolve on their data as sister group to Branchiura, and not as ingroup fish lice. Indeed a taxon name Ichthyostraca sensu Zrzavý, 2001 has been proposed for the putative (Pentastomida þ Branchiura) clade. The implication would be that pentastomids are some sort of fish louse-like crustacean, presumably carried onto land in the bodies of the early terrestrial vertebrates. Martin & Davis (2001) accepted Pentastomida as ingroup Crustacea in their authoratiative summary of crustacean systematics and other modern summaries of crustacean phylogeny have done likewise (e.g. Richter et al. 2009). By contrast, Walossek & Mçller (1994), Walozsek et al. (2006) and Almeida et al. (2008a) interpreted pentastomids as part of the arthropod stem group, a model closer to the Heymons concept and supported by the fact that pentastomids seem to lack obvious morphological apomorphies characterising Euarthropoda and/or crown-group Crustacea. In a critique of the Ichthyostraca hypothesis Walossek & Mçller (1994) and Walozsek et al. (2006) noted that other parasitic crustaceans can still be placed quite easily into their appropriate groups based at least on larval characters. They also noted differences in developmental biology, tagmosis and limb structure which, for them, ruled out euarthropod affinities; see these authors for details. Almeida et al. s (2008b) morphology-based cladogram resolved pentastomids among the Ecdyzoa. Specifically, pentastomids emerged somewhat controversially in a position between the tardigrades and various nematohelminth worms (nematodes, priapulids, kinorhynchs, etc). These authors also attempted to integrate pentastomids into phylogenies of early fossil lobopodian/arthropod taxa; although their study could be criticised for scoring all euarthropods as a single terminal Arthropoda and thus not offering an explicit test of the Branchiura/Crustacea hypothesis. Further work is evidently needed to resolve pentatstomid affinities. Host parasite evolution Irrespective of their relationships, it is interesting to speculate on the definitive hosts of pentastomids in the Cambrian (Walossek & Mçller 1994), before true vertebrates had become properly established. Conodonts, as putatative stem-chordates with a Cambrian fossil record, have museum-zoosyst.evol.wiley-vch.de

5 Zoosyst. Evol. 86 (1) 2010, sometimes been mentioned in this context (e.g. Schram in Martin & Davis 2001). In general, pentastomids are one of the few parasitic groups which inhabit the vertebrate respiratory system as adults as opposed to the gut or bloodstream and it is conceivable that films of water in the lungs are a substitute for their original aquatic environments. Likewise, we must consider what their hosts were during the (Devonian) period of terrestrialisation through to the (Carboniferous) origin of the reptiles. Although some modern pentastomid genera such as Leiperia Sambon, 1922 use (freshwater) fish as intermediate hosts, there is no documented evidence of marine pentastomids retained in a putative definitive fish host. Various amphibians can act as intermediate hosts (Norval et al. 2009) and they have also recently been identified as definitive hosts too (e.g. Barton & Riley 2004). Condensing the phylogenies proposed by Almeida & Christoffersen (1999) and Waloszek et al. (2006: fig. 6), there seems to be basic pattern of (Palaeozoic stem taxa (cephalobaeinids (raillietiellids (reighardiids (lingulatoids þ porocephaloids))))). This shows little phylogenetic correlation with these animals respective definitive hosts. Basal pentastomids are not obviously characteristic for basal reptile lineages, occurring in snakes and even marsupials (Spratt 2003). The unusual Reighardia Ward, 1899 has a highly specialised host relationship with seabirds, with whom they seem to practice autoinfection (see below) in which case there is no intermediate host. Only Diesengia Sambon, 1922 and Pelonia Junker & Boomker, 2002 appear to be associated with a primitive reptile group (the chelonians: i.e. tortoises, terrapins and turtles), but these pentastomid genera resolve among the derived porocephaloids. The original (definitive) host of the modern pentastomids is difficult to reconstruct. It would be tempting to infer a snake the oldest fossils of which are currently late Cretaceous in age (ApesteguÌa & Zaher 2006) although some authors consider snakes to be derived from varanoids (monitor lizards), which also act as definitive hosts for the porocephaloid genus Samboina Noc & Giglioli, 1922 among the modern pentastomids. Material and methods Data were drawn from available catalogues and original labels, crossreferenced against the literature. In particular Heymons twenty-three publications on the subject were carefully checked for the descriptions of new taxa (Tab. 1); to see which of them should be associated with type material in Berlin. With the exception of some specimens loaned in the late 1970s catalogue numbers had, in the main, not been previously assigned. Thus during the course of this project each unregistered lot was assigned a number under the traditional acronym ZMB/ Arach (Zoologisches Museum Berlin). The data gathered here will be entered into the SysTax database ( and will eventually be mirrored into the museum s own database using the platform Specify. Identifications of non-type material could not be checked and they are listed here under the names given on the specimen labels; with a couple of exceptions discussed under the Remarks to each species. The systematic structure and sequence of family-group names adopted here broadly follows Almeida & Christoffersen (1999). In the collection itself, type specimens have been separated from the non-type material and these non-types have been arranged in the same phylogenetic sequence as the one presented here. Correct current names for reptilian hosts were checked against the TIGR Reptile Database ( amphibian hosts were checked against the database Amphibian Species of the World 5.3 ( and intermediate fish hosts were checked against FishBase ( those of other taxa against various library and online resources. Following the recommendations of the Standard Nomenclature of Parasitic Diseases (SNOPAD) (see e.g. Kassai et al. 1988) the terms pentastomosis, linguatulosis and porocephalosis are used when refering to pentastomid infection. One of us (JHG) also examined material collected by Natterer in the Naturhistorisches Museum Wien, which presumably includes the specimens described by Diesing (see above). Historically, Berlin s Pentastomida were associated with the arachnid and myriapod collection, possibly because of earlier hypotheses of affinities with mites or myriapods and probably also because the stem-arthropod groups Onychophora and Tardigrada are held under this curatorship too. Given the uncertainties about the affinities of pentastomids (see above) there are, at present, no plans to transfer them to the crustacean curatorship. Type collection affinis Bovien, 1927 Figure 2l Raillietiella affinis Bovien, 1927: 5 6, figs 4 6. Type locality. [Jombang], [East] Java, Indonesia. Host. The tokay gecko Gekko gecko Linnaeus, 1758 [as the junior synonym Gekko verticillatus Laurenti, 1768] (Sauria: Gekkonidae) or Panther-Gecko. Type material. Syntype (as cotype ), ZMB 48037; 3 specimens; [leg. P. Bovien, 1923]; ex Coll. Heymons nr. 29. Present name. Raillietiella affinis Bovien, Remarks. According to Ali et al. (1982a), Bovien s type series an immature slide-mounted female and two smaller cotype females in alcohol is deposited in the Natural History Museum, London. They did not mention a Berlin (alcohol) specimen. We can only assume that our example was donated to, or exchanged with, Heymons (see Richard Heymons), which would explain why it is associated with one of his collection numbers. We can be confident that this is part of the type series given the cotype label and the fact that the host data matches the original description. We can also narrow down the locality based on the original paper to Jombang [as Djombang ] in East Java where Bovien worked from at the Java Sugar Experiment Station (Pasoeroean). Hett (1934) suggested that Raillietiella affinis Bovien, 1927 was a junior synonym of R. indica Gedoelst, This was disputed by Ali et al. (1982a), who maintained R. affinis as a separate species; see also Ali et al. (1985). bicaudata Heymons & Vitzthum, 1935 Figure 2e Raillietiella bicaudata Heymons & Vitzthum, 1935: 21 23, fig. 15. Type locality. Berlin Aquarium. museum-zoosyst.evol.wiley-vch.de

6 134 Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Host. According to the original description by Heymons & Vitzthum, the type examples came from the lungs of a common kingsnake Lampropeltis [as Ophibolus] getula Linnaeus, 1766 (Serpentes: Colubridae) in the Berlin aquarium. We were unable to trace material with exactly this provenance. The authors also explicitly mentioned material from Elaphe couperi presumably the eastern indigo snake Drymarchon couperi Holbrook, 1842 (Serpentes: Colubridae) for which a matching label could be found. However, another label mentions the western indigo snake Drymarchon [as Spilotes] corais Boie, 1827 (again Serpentes: Colubridae). Figure 2. Overview of the Pentastomida type specimens in alcohol in the Museum fçr Naturkunde Berlin; showing the condition of the specimens and their relevent labels. a b. Raillietella kochi Heymons, 1926, syntypes, ZMB 17165, c. Raillietella shipleyi Heymons, 1926 [= R. kochi], syntypes, ZMB 17166, d. Raillietiella mabuiae Heymons, 1922, syntypes, ZMB 17167, e. Raillietiella bicaudata Heymons & Vitzthum, 1935, syntype?, ZMB 17168, f h. Cephalobaena tetrapoda Heymons, 1922, syntypes, ZMB , i j. Pentastomum javanicum Bovien, 1927 [= Kiricephalus pattoni], cotypes ZMB , k. Elenia lialisi Heymons, 1939 [= Waddycephalus sp.], cotype ZMB 48024, l. Raillietiella affinis Bovien, 1927, cotype ZMB museum-zoosyst.evol.wiley-vch.de

7 Zoosyst. Evol. 86 (1) 2010, Type material. Syntype, ZMB 17168; collector not recorded, 03.V.[19]23; ex Coll. Heymons, nr. 63 (the couperi specimen). Present name. Raillietiella bicaudata Heymons & Vitzthum, Remarks. Both the original description and the present data on the labels are rather confusing, and it is not clear whether the Berlin specimen is really part of the original type series. According to available data two specimens should be present, but only one could be found in the jar. At least one label is obscure in its meaning in V...ing Koch (No. 9) m... R. bicaudata angetroffen R. Heymons This would postdate the year of publication and implies non-type specimens. Since no further indication of an alternative type repository other than Berlin is given by Heymons & Vitzthum (1935) we assume that the specimens retained here could be part of the type series. javanicum Bovien, 1927 Figures 2i j Pentastomum javanicum Bovien, 1927: 2 4, figs 1 4. Type locality. [Jombang], [East] Java, Indonesia. Host. The crab-eating frog Fejervarya [as Rana] cancrivora (Gravenhorst, 1829) (Anura: Ranidae) or Philippinen-Frosch. Type material. Syntypes (as cotypes ), ZMB 48022; 4 juveniles; leg. P. Bovien, [1923]. Syntype (as cotype ) ZMB 48023; 1 juvenile; leg. P. Bovien, [1923]; ex Coll. Heymons nr. 54. Present name. Junior synonym of Kiricephalus pattoni (Stephens, 1908). Remarks. As with R. affinis, part of Bovien s type material seems to have been sent to Heymons. Again, the material is explicitly labelled as cotype and thus data from the original description can be used to augment the locality and date of collection. One point of conflict is that the collection data gives the host as Fejervarya cancrivora while the original description states Rana tigrina [now Hoplobatrachus tigerinus (Daudin, 1802)], the Indus Valley Bullfrog. Pentastomum javanicum was based on juveniles only. Heymons (1935: 168) mentions Pentastomum javanicum in Rana cancrivora (= Rana tigrina), but confusingly javanicum is not listed either as a valid species or as a synonym in his systematic section of the same work, or in the systematic index. The closest we can come to resolving the issue is to note that under Kiricephalus pattoni there is a statement Heymons (1935: 235) Außerdem sind Larven in javanischen Fræschen, Rana cancrivora (tigrina) beobachtet worden (Bovien, Sambon) [Additionally larvae have been observed in the Javan frog Rana cancrivora (tigrina) (Bovien, Sambon)]. The label on our museum jar has also been corrected by hand at some stage to = Kiricephalus pattoni and thus we presume Bovien s species to be a synonym; even though we have not been able to trace the author who formally proposed this. kochi Heymons, 1926 Figures 2a b, 3a b Raillietella kochi Heymons, 1926a: 45 46, figs 1 2. Type locality. Berlin Aquarium, Ethiopia [as Abessienien ] ¼ patria host. In the original description Heymons refers to an abessynischen Warans, but it is unclear if the host animal was wild-caught before being brought to Berlin. Host. From both lungs of a savannah monitor lizard Varanus exanthematicus Bosc, 1792 [as its synonym V. ocellatus Heyden in Rçppel, 1830] (Sauria Varanidae) or Steppenwaran. Type material. Syntypes, ZMB 17165; 1 < (damaged), 3,; collector not recorded, 14.VIII.1925; Presumably part of the original Heymons series nr. 24 (see below); alcohol and slide material. Note that the three females were inadvertently labelled ZMB (cf. R. shipleyi) (Figs 2a c). Present name. Raillietella kochi Heymons, Remarks. There are also two microscope slides, both dissected from syntype specimens with V. ocellatus as the host and both numbered ZMB One is labeled posterior hook of female, the other male copulatory spicule. These slides were almost certainly made by John Riley who borrowed the specimens in 1979; see also Ali et al. (1985). A paratype of this species was also deposited in the Hamburg Museum (Weidner 1959: 93). lialisi Heymons, 1939a Figure 2k Elenia lialisi Heymons, 1939a: , figs 3 4. Type locality. Iffar, Dutsch New Guinea [Indonesia as Dutch New Guinea ]. Host. The legless lizard or flop-foot Lialis jicari Boulenger, 1903 (Reptilia: Pygopodidae) or Neuguinea-Flossenfuß Type material. Syntype (as cotype ) ZMB 48024; numerous juveniles; collector not recorded; ex Coll. Heymons nr. 2. Present name. Waddycephalus sp.; effectively a nomen dubium. Remarks. The original label bears the comment erhalten vom British Museum London 1939, (in litt.) larvae [received from the British Museum London 1939, (in press) larva]. It is not clear from the rather equivocal original spelling on the label whether the locality data refers to Dutch New Guinea (now part of Indonesia) or Deutsch New Guinea (which would fall under modern Papua New Guinea). Iffar, however, belongs to the Indonesian side of the island. This species was not listed by Self (1969), and Riley et al. (1985: 47), in their review of Australasian pentastomids, explicitly stated that: On the available evidence, Heymon s nymphs from L. jicari deserve only tentative generic status and do not warrant specific status. These authors also treated the type locality as Indonesian rather than Papuan in origin. museum-zoosyst.evol.wiley-vch.de

8 136 Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Type material. 3 syntypes, ZMB 17167; leg. L. Schulze, VI.1903; label bears the inscriptions No. 70, No. 18 and Formol 4 % ; alcohol and microscope slide. Present name. Raillietiella mabuiae Heymons, Remarks. There is also one microscope slide, dissected from a syntype specimen with Mabuya sulcata as the host and numbered ZMB It is labeled male copulatory spicule. The slide was almost certainly made by John Riley who borrowed the specimens in 1979; see also Ali et al. (1985). shipleyi Heymons, 1926 Figure 2c, 3d e Raillietella shipleyi Heymons, 1926a: 46 47, fig. 3. Type locality. Not recorded. Host. From the lung of a savannah monitor lizard label states both Varanus exanthematicus Bosc, 1792 [as its synonym V. ocellatus Heyden in Rçppel, 1830] (Sauria: Varanidae) or Steppenwaran. Type material. 3 Syntypes, ZMB 17166; collector and date not recorded; ex Coll. Heymons nr. 28; alcohol and microscope slides. Present name. Junior synonym of Raillietiella kochi Heymons, 1926, which originates from the same host; synonymised by Ali et al. (1985: 114). Remarks. There are also two microscope slides, both dissected from syntype specimens with Varanus exanthematicus as the host and both numbered ZMB One is labeled posterior hook of,, the other male copulatory spicule. The slides were almost certainly made by John Riley who borrowed the specimens in 1979; see also Ali et al. (1985). tetrapoda Heymons, 1922 Figure 2f h Cephalobaena tetrapoda Heymons, 1922: , fig. 4. Type locality. San Bernadino, Departamento de Cordillera, Paraguay. Host. From the Neotropical pit viper genus Lachesis sp. (Serpentes: Viperidae) or Buschmeisterschlange. Type material. Syntypes ZMB 48019, 2,; leg. R. Fiebrig, 24.IV[19]06; also bears label F.Nr.: 373/06. Figure 3. Slide mounted type and non-type pentatstomid material in the Museum fçr Naturkunde Berlin. a b. Raillietella kochi Heymons, 1926, syntypes, ZMB 17165, c. Raillietiella mabuiae Heymons, 1922, syntypes, ZMB 17167, d e. Raillietella shipleyi Heymons, 1926 [= R. kochi], syntypes, ZMB (all dissected and prepared by John Riley), f g. examples of historical whole-animal mounts of unidentified pentastomids. mabuiae Heymons, 1922 Figures 2d, 3c Raillietiella mabuiae Heymons, 1922: , fig. 3. Type locality. Namibia, as Deutsch-Sçdwest-Africa. Host. From the lung of the western rock skink Trachylepis [as Mabuia] sulcata (Peters, 1867) (Sauria: Scincidae) or Felsenskink. Syntypes ZMB 48020, 3 <; leg. R. Fiebrig, 24.IV.[19]06; also bears label F.Nr.: 373/06. Syntypes ZMB 48021, 1 <; leg. R. Fiebrig, 24.IV.[19]06; also bears label F.Nr.: 373/06. Present name. Cephalobaena tetrapoda Heymons, Lost type? travassosi Heymons, 1932 Alofia travassosi Heymons, 1932b: , figs 1 2. Type locality. Basey River, Samar Island, Phillippines. Host. Not recorded. museum-zoosyst.evol.wiley-vch.de

9 Zoosyst. Evol. 86 (1) 2010, Material. 2,; leg. Jagor; the repository was explicitly stated in the description by Heymons as Zoologisches Museum Berlin. Material matching this data could not be traced in the collection during the preparation of this paper. Present name. Elenia travassosi (Heymons, 1932b). Unpublished type One named specimen (ZMB 48054); ex. Coll. Heymons nr. 21 bears the inscription aus dem Typenmaterial von Lohrmann ; including host data relating to a purple swamphen Porphyrio sp. (Aves: Rallidae) or Purpurhuhn. A further note reidentifies it as Armillifer armillatus. As far as we are aware Lohrmann published only one paper on pentastomids (Lohrmann 1889) in which he created two new species. However, neither of these matches the name on the ZMB label and we suspect that the Berlin specimen refers to an unpublished taxon which we will not mention here to avoid creating a nomen nudum. In the same paper Lohrman refers to... endlich eine Jugendform mit wulstig aufgetriebenen Leibesringen, von der ich nicht zweifle, dass sie zu P. multicinctum, Harl. gehært, und die eingekapselt in einem Purpurhuhn (Porphyrio) gefunden worden war. [... finally a young form with bulging body rings which I don t doubt belongs to P. multicinctum Harl. and which was found encapsulated in a purple swamphen (Pophyrio)]. The species Pentastomum multicinctum Harley, 1857 is a junior synonym of Cubirea [formerly Armillifer] annulatus (Baird, 1853). It is concievable that this note from Lohrmann refers to our material and that at some stage he (or someone?) considered raising a new taxon for it. Its presence in a bird is unusual and was commented upon by Heymons (1935: 240) again with specific reference to Lohrmann s swamphen specimens although as an encysted larvae this presumably refers to an intermediate host. Non type collection Class Pentastomida Diesing, 1835 Remarks. Traditionally, Pentastomida has been treated as a class (or even a phylum) comprising two orders: Cephalobaenida and Porocephalida. The name Eupentastomida Waloszek et al., 2006 was introduced for all extant pentastomids, to the exclusion of the fossil taxa. The higher systematics of the extant tongue worms are frustrating, in that there are at least two alternative schemes available in the literature. In their summary of crustacean higher taxa Martill & Davis (2001) recognised an order Cephalobaenida (Families: Cephalobaenidae and Reighardiidae) and an order Porocephalida (Families: Armilliferidae, Diesingidae, Linguatulidae, Porocephalidae, Sambonidae, Sebekiidae and Subtriquetridae). Oddly, they ascribe authorship of many of these families to Fain (1961), despite the fact that the names in question were clearly introduced as family or subfamily groups by earlier workers (cf. Heymons 1935; Self 1969). By contrast the cladistic-based analysis of Almeida and Christofferson (1999) recognised four orders Cepalobaenida (Family: Cephalobanidae), Raillietellida (Family: Raillietellidae), Reighardiida (Family: Reighardiidae) and Porocephalida. The latter comprised a superfamily Lingulatoidea (Families: Linguatulidae and Subtriquetridae) and a superfamily Porocephaloidea (Families: Porocephalidae and Sebekidae). However, at least the names Raillietellida and Reighardiida have not become established in the subsequent literature (Riley et al. 2003; Spratt 2003; Abreu-Acosta et al. 2006) and, for example, standard databases such as the World Register of Marine Species (WoRMS) continue to cite Reighardia among the cephalobaeinids. In the absence of an authoritative modern catalogue of tongue worms the last attempt to list all taxa we are aware of was by Self (1969) the Almeida and Christofferson scheme is tentatively adopted here as offering some basic phylogenetic structure. Order Cephalobaenida Heymons, 1935 Family Cephalobaenidae Heymons, 1935 Cephalobaena Heymons, 1922 Cephalobaena tetrapoda Heymons, 1922 Material. ZMB 48030, 1 specimen; Argentina; leg. D. L. Cernosvitov, date and host not recorded [but see below]; ex Coll. Heymons nr. 30. ZMB 48031, 3 specimens (1 damaged); Loreto near Santa Ana, Misiones Province, Argentina; leg. D. L. Cernosvitov, 16.IX.1932; from the lungs of a racer Leptophis ahaetulla ahaetulla (Linnaeus, 1758) [as the junior synonym Leptophis liocercus (Wied, 1824)] (Serpentes: Colubridae) or Schlanknatter ; ex Coll. Heymons nr. 30. ZMB 48032, 4 specimens; Loreto near Santa Ana, Misiones Province, Argentina; leg. D. A. Ogloblin, 1931; host not recorded; ex Coll. Heymons nr. 30. ZMB 48035, 3 specimens; Argentina; collector not recorded; from the tropical pit viper genus Lachesis (Serpentes: Viperidae) or Buschmeisterschlange ; ex Coll. Heymons nr. 31. Remarks. This unusual-looking species with its more elongate cephalic legs and a mouth on a similar blunt protuberance is widely regarded as the most primitive of the living pentastomids (cf. Almeida & Christoffersen 1999). Details of its morphology and development can be found in, e.g., von Haffner & Rack (1971) and recent accounts of this species and its snake hosts can be found in Almeida et al. (2006, 2008b). The Argentinean specimens listed above do not belong to the type series (see above) since they (largely) postdate the species description, which is also restricted in the original diagnosis to the material from Paraguay. Order Raillietiellida Almeida & Christoffersen, 1999 Family Railietiellidae Sambon, 1922 Raillietiella Sambon, 1922 Raillietiella boulengeri (Vaney & Sambon, 1910) Figure 4a Material. ZMB 48034; 1 <; Berlin Aquarium; collector not recorded, 21.VII.[19]31; from the Gabon viper Bitis gabonica Dum ril, Bibron museum-zoosyst.evol.wiley-vch.de

10 138 Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Figure 4. Examples of non-type pentastomids in the Berlin collection. a. Raillietiella boulengeri (Vaney & Sambon, 1910), ZMB 48043, b. Raillietiella orientalis (Hett, 1915), ZMB 48038, c. Linguatula serrata Frælich, 1789, ZMB 48141, d. Armillifer armillatus (Wyman, 1847) ZMB 48056, e. Armillifer mazzai (Sambon, 1922), ZMB 48178, f. Cubirea pomeroyi (Woodland, 1921), ZMB & Dum ril, 1854 (Serpentes: Viperidae) or Gabunviper ; ex Coll. Heymons nr. 22. ZMB 48040, 2,; tropical Africa ; collector not recorded, 07.XI.1910; from the lung of a puff adder Bitis sp. (Serpentes: Viperidae) or Puffotter ; label bears the inscription Porocephalus armillatus, bestimmt von Prof. von Linstow, Zoologisches Museum. ZMB 48042, 1 specimen; Berlin Aquarium; collector not recorded, 21.VII.1931; from the Gaboon viper Bitis gabonica (Serpentes: Viperidae) or Gabunviper ; ex Coll. Heymons nr. 23. museum-zoosyst.evol.wiley-vch.de

11 Zoosyst. Evol. 86 (1) 2010, ZMB 48043, 3 specimens; Berlin Aquarium; collector not recorded, 21.VII.[19]31; from the Gaboon viper Bitis gabonica (Serpentes: Viperidae) or Gabunviper ; ex Coll. Heymons nr. 23. ZMB 48044, 1 specimen; Berlin Aquarium; collector not recorded, 20.I.[19]32; from the lungs of a Bastard (hybrid) of the water snake Nerodia [as Tropidonotus] fasciata Linneaus, 1766 and another species (whose name as we read it does not appear in any databases) (Serpentes: Colubridae) or einer Schwimmnatter und eine...? ; ex Coll. Heymons nr. 34. Remarks. Ali et al. (1982b) confirmed the validity of this species as part of a detailed redescription. Raillietiella furcocera (Diesing, 1835) Material. ZMB 48025; 1 specimen; San Bernadino, Paraguay; leg. Fiebrig, 27.VII.[19]05; from a jarrara Lachesis sp. (Serpentes: Viperidae) or Buschmeisterschlange ; label bears the inscriptions Fiebrig Nr and S ; 1 specimen; San Bernadino, Paraguay; leg. Fiebrig, 27.VII.05; from the woodland snake Rhadinaea furca [species name appears to be incorrect; it could not be traced in The Reptile Database] (Serpentes: Colubridae) or Yarrara ; label bears the inscription Fiebrig No Remarks. There seems to be some confusion concerning the hosts for this species, given that Lachesis has not been recorded from Paraguay and that the common names jarrara/yarrara are here applied to two different genera of snake. The names probably refer to Bothrops jararaca Wied, 1824 (Serpentes: Viperidae) which is known from Paraguay. Raillietiella hemidactyli Hett, 1934 Material. ZMB 48027; 1 specimen labelled as Raillietiella geckonis (Diesing, 1850); India; collector and date not recorded; from the Oriental garden lizard Calotes versicolor (Daudin, 1802) (Sauria: Agamidae) or Blutsaugeragame ; ex Coll. Heymons nr. 4; label bears the inscriptions erhalten vom British Museum London 1939, det. R. Heymons and S 3. ZMB 48028; 1 specimen labelled as Raillietiella geckonis; India; collector and date not recorded; from the Oriental garden lizard Calotes versicolor (Sauria: Agamidae) or Blutsaugeragame ; ex Coll. Heymons nr. 4; label bears the inscription erhalten vom British Museum London 1939, det. R. Heymons and S 3. Remarks. Ali et al. (1982a) discussed the problems associated with the species name Raillietiella geckonis, including the unsatisfactory original description and confusion about the true host species of the type material. They concluded that the name should be abandoned. Interestingly, they explicitly referred to some Indian material in the London Natural History Museum recovered from the garden lizard Calotes versicolor which was identified by Richard Heymons as R. geckonis, but which Ali et al. (1981) had previously reidentified as Raillietiella hemidactyli Hett, The Berlin specimens were not explicitly mentioned by Ali et al. (1981, 1982a), but clearly share the same locality and host data, originate from London and were expliticly identified by Heymons as R. geckonis: det R. Heymons. We know that there was an exchange of material between Heymons and London and thus we strongly suspect that ZMB are part of the same series, the London examples of which were reidentified by Ali et al. (1981). For this reason we reassign these Berlin specimens to R. hemidactyli as opposed to R. geckonis; which is in any case a putative nomen dubium. Raillietiella mediterenea (Hett, 1915) Material. ZMB 48029; 2 specimens; Berlin Aquarium; collector not recorded, 14.VII.1925; from the lung of Dahl s wipe snake Platyceps najadum Eichwald, 1831 [as the junior synonym Coluber [Zamenis] dahlii (Fitzinger, 1826)] or Schlanknatter ; ex Coll. Heymons nr. 27. Remarks. Listed as a valid species by Ali et al. (1985). Raillietiella orientalis (Hett, 1915) Figure 4b Material. ZMB 48036; 1 specimen; [?Berlin] Aquarium; collector not recorded, 29.VI.1925; from a female striped aesculapian rat snake Zamenis [as Elaphe] lineatus (Serpentes: Colubridae) or Italienische Øskulapnatter ; patria host Italy; ex Coll. Heymons nr. 32, host may actually be Zamenis longissimus Laurenti, ZMB 48038; 1,; Berlin Aquarium; collector not recorded, 31.III.1937; from the oriental rat snake Ptyas mucosa [as Coluber (Zamenis) mucosa] (Serpentes: Colubridae) or Asiatische Rattenschlange ; ex Coll. Heymons nr. 76. ZMB 48039; 1 <; Berlin Aquarium; collector not recorded, 31.III.1937; from the oriental rattlesnake Ptyas mucosa Linnaeus, 1758 [as Coluber (Zamenis) mucosa] (Serpentes: Colubridae) or Asiatische Rattenschlange ; ex Coll. Heymons nr. 76. ZMB 48041; 3 specimens; India, leg. Ræssner, III.1911; from the lung of a cobra Naja sp. (Serpentes: Elapidae) or Kobra ; ex Coll. Heymons nr. 25, label bears the inscription S 8. ZMB 48045; numerous specimens;?india (see above), leg. Ræssner, III.1911; from the lung of a cobra Naja sp. (Serpentes: Elapidae) or Kobra ; original label bears the inscription Naja spec. mit ungekapselten Larven in der Leber, Porocephalus and Nr. 17a [Naja spec. with unhatched larvae in the liver...], presumably originally part of the series Heymons nr. 25. ZMB 48046; 10 specimens (4 damaged);?india (see above), leg. Ræssner, III.1911; from the lung of a cobra Naja sp. (Serpentes: Elapidae) or Kobra ; original label bears the inscription Naja spec. mit ungekapselten Larven in der Leber, Porocephalus and Nr 17a [Naja spec. with unhatched larvae in the liver...], presumably originally part of the series Heymons nr. 25. Remarks. Ali et al. (1982b) confirmed the validity of this species in a detailed redescription. Raillietiella sp. Material. ZMB 48033, numerous specimens; 2 Meilen æstl. Algier [2 miles east of Algiers]; Algeria; leg. V. Gotsek; date and host not recorded. museum-zoosyst.evol.wiley-vch.de

12 140 Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Order Reighardiida Almeida & Christoffersen, 1999 Family Reighardiidae Heymons & Vitzthum, 1936 Reighardia Ward, 1899 Reighardia sternae (Diesing, 1864) Material. ZMB 48047; 1, specimen; Rybatschi [as Rossitten ], near Kaliningrad, Baltic Coast of Russia; collector not recorded, VIII.1935; from the Arctic tern Sterna paradisaea Pontoppidan, 1763 [as S. macrura] (Aves: Sternidae) or Kçstenseeschwalbe ; ex Coll. Heymons nr. 58, label bears the inscription von Dr. L. Szidat (Rossitten) 1935 erhalten [recieved from]. ZMB 48048; 2 specimens, damaged by mould; northern Greenland; collector and date not recorded but from the Zoological Museum Copenhagen; from a Mæwe [seagull] (Aves: Laridae); ex Coll. Heymons nr. 57, label bears the inscription im Austausch vom Zoologischen Museum Kopenhagen erhalten R. H. Remarks. Until recently, Reighardia was the only pentastomid genus known to habitually parasitize birds a further species, Reighardia lomviae Dyk, 1975, has been described from the guillemot. Since this time a second reighardiid species has been described in a new genus from Spanish vultures (MartÌnez et al. 2004), while a Raillietiella species has been found in vultures from Pakistan (Riley et al. 2003). By contrast Reighardia sternae occurs in gulls and other sea birds (Riley 1972; Bakke 1972) and there is good evidence that avian pentastomids are transferred directly and/or autoreinfect the host (e.g. Banaja et al. 1976; Bæckler 1984), without the need for an intermediate host taxon. The morphology of R. sternae has been described in detail by Riley (1973a, b). The record from as far north as Greenland is not unreasonable. Sprehn (1928) recorded both this species, and Linguatula serrata, from the Arctic region. Order Porocephalida Heymons, 1935 Superfamily Linguatuloidea Leuckart, 1860 Family Linguatulidae Leuckart, 1860 Linguatula Frælich, 1789 Linguatula recurvata (Diesing, 1850) Material. ZMB 48139; 1 juvenile; Surinam [as Dutch Guyana ]; collector and date not recorded; from the endocardium of a white-lipped peccary Tayassu pecari (Fischer, 1814) [= Dicotyles labiatus] (Mammalia: Tayassuidae) or Weißbartpekari ; label bears the inscription 13. r. Remarks. Recent studies maintain Linguatula recurvata as a species distinct from the better known, and medically significant, L. serrata (see below); see e.g. Arroyo et al. (1986) for a differential diagnosis. There is so far no report of L. recurvata affecting humans. Linguatula serrata Frælich, 1789 Figure 4c Material. ZMB 48140; numerous specimens; Berlin Zoological Garden; collector not recorded, 21.I.[19]23; from the lymph nodes in the mesenterium of a nilgai Boselaphus tragocamelus (Pallas, 1766) (Mammalia: Bovidae) or Nilgau-Antilope ; label bears the inscription Zoologischer Garten Berlin, Zoologisches Museum Berlin, E.K.N ZMB 48141; 5 specimens originally from the teaching collection Schausammlung Studenten ; locality, collector and date not recorded; from the sinufrontal of a domestic dog Canis [lupus] familiaris (Linnaeus, 1758) or Haushund ; label bears the inscription Zoologisches Museum Berlin 737. ZMB 48142; 1 specimen; Sri Lanka [as Ceylon ], leg. Nietner, date and host not recorded; label bears the older number ZMB 48143; 2 specimens; Berlin Zoological Garden; collector and date not recorded; from the lung of a goitered gazelle Gazella subgutturosa Gçldenstådt, 1780 (Mammalia: Bovidae) or Kropfgazelle. ZMB 48144; 10 otherwise unlabelled specimens; locality, collector, date and host not recorded. ZMB 48145; 15 specimens; from Hejaz [as Hedschas ], Saudia Arabia via the Berlin Zoological Garden; collector and date not recorded; from the lung of an east African oryx Oryx beisa (Rçppel, 1835) (Mammalia: Bovidae) or Oryxantilope ; label bears the inscription... vor 4 Wochen çber Jaffa aus Hedschas eingefçhrt. Hier im Zoologischen Garten eingegangen. ZMB 48146; 18 specimens; locality, collector, date and host not recorded, but probably from the lymph glands of a domestic goat Capra aegagrus hircus Linnaeus, 1758 (Mammalia: Bovidae) or Hausziege ; label bears the inscription No. 10 and Gland. Lymph, Entozoa hircus. ZMB ; numerous specimens, in part from teaching collection Schausammlung Studenten, in part dried out; locality, collector, date and host not recorded throughout. ZMB 48154; 1 specimen; locality, collector and date not recorded; from a domestic dog Canis [lupus] familiaris (Mammalia: Canidae) or Haushund ; label bears the inscription Zoologisches Museum Berlin 798. Remarks. As implied by the collection data, both wild (Sachs et al. 1973) and domestic ungulates are common intermediate hosts. Dogs, and other carnivores, are the final host; hence the occasional referral to these animals as dog tongueworms. Linguatula serrata is a widespread species, particularly in the tropics and a summary of its biology can be found in Heymons (1942). It can also affect humans (Fain 1975) see also Human infection both as intermediate (visceral) and as final (nasopharyngeal) hosts. Note that the author s name is frequently misspelled Fræhlich or Frohlich in the literature. Superfamily Porocephaloidea Sambon, 1922 Family Porocephalidae Sambon, 1922 Armillifer Sambon, 1922 Armillifer armillatus (Wyman, 1847) Figure 4d Material. ZMB 4180; 1 specimen; Africa, collector and date not recorded; from the abdominal cavity of a chimpanzee Troglodytes niger [now Pan troglodytes (Blumenbach, 1775) (Mammalia: Hominidae) or Schimpanse. ZMB 48056; 2 specimens; Africa; collector and date not recorded; from either the royal (Python regius (Shaw, 1802)) or rock (P. sebae (Gmelin, 1789)) python (Serpentes: Pythonidae) or Kænigspython oder Nærdliche Felsenpython ; label bears the inscriptions Heymons det., I., Ab. B. and 5.. museum-zoosyst.evol.wiley-vch.de

13 Zoosyst. Evol. 86 (1) 2010, ZMB 48057; 1 <, 1 juvenile,; Rutshuru, Democratic Republic of the Congo [as Belgisch-Kongo ]; collector and date not recorded; from the viper Vipera sp. (Serpentes: Viperidae) or Viper ; ex Coll. Heymons nr. 68. ZMB 48058; 3 juveniles; Rutshuru, Democratic Republic of the Congo [as Belgisch-Kongo ]; collector and date not recorded; aus Viverra spec. or Schleichkatzen. The putative host must be treated with caution since civets (Viverra sp.) are typically south-east Asian in their distribution. Maybe a misinterpretation of Vipera, as per the previous ZMB number from the same locality? ZMB 48059; 1 <, 1,?, juvenile,; Tesa, Uganda; collector and date not recorded; from a hedgehog or Igel (Mammalia: Erinaceidae); received from the British Museum London and bearing the labels Heymons det. and No. 3. ZMB 48060; 2 specimens; Africa; collector not recorded, 22.V.[18]90; host not recorded; one label illegible. ZMB 48061; many larvae plus larvae with the organs of the host; Tirko, Cameroon; collector and date not recorded; probably from the duiker Cephalophus sp. (Mammalia: Bovidae) or Schopfducker. ZMB 48062; larvae on the spleen of the host; Berlin Zoological Garden; leg. Collin, date not recorded; from a male serval Felis [now Leptailurus] serval (Schreber, 1776) (Mammalia: Felidae) or Serval, patria host Senegambia, Africa. ZMB 48063; larvae with the organs of the host; Biologisches Landwirtschaftliches Institut Amani in Ostafrika, Tanzania; collector, date and host not recorded; label bears the inscription Jr. No. 644/09, Zoologisches Museum Berlin, Jr No. 644/09. ZMB 48064; many larvae in connective tissue, 1 < and 1, extra in jar; locality, collector and date not recorded; from a female baboon (Mammalia: Cercopithecidae) or Pavian ; label bears the inscription Zoologisches Museum Berlin, E.K.N ZMB 48065; many larvae with the organs of the host; Berlin Zoological Garden; leg. Collin, 30.IV.[18]92; from a serval Felis [now Leptailurus] serval (Mammalia: Felidae) or Serval. ZMB 48066; larvae; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscriptions i. d. Fundortliste unter Nummer 14 and Zoologisches Museum Berlin Jr. No. 604/1910, Zoolog. Museum Berlin. ZMB 48067; larvae; Cameroon; leg. Schåfer, date not recorded; label bears the inscription Fundort i. d. Fundortliste unter Nummer 26 and Zoologisches Museum Berlin, Jr. No. 1168/10. ZMB 48068; 1 specimen; locality, collector and date not recorded; from a python lung Python sp. (Serpentes: Pythonidae) or Python ; the label bears the inscription Coll. Buxton No. 2 and bears what looks like a subspecies name, but one for which we could find no record in the available literature. There is no indication this should be the type of a new subspecies. ZMB 48069; 2 specimens with a piece of the host s liver; Cameroon; leg. Schåfer, date not recorded; from the duiker Cephalophus sp. (Mammalia: Bovidae) or Antilope ; label bears the Fundort s. Liste Nr. 14 and Zoologisches Museum Berlin, Jr. No. 1168/10. ZMB 48070; 4 specimens with the organs of the host; Bismarkburg, Togo; collector and host not recorded, 14.V[18]93; label bears the inscription No. 3 and No. 82. ZMB 48071; 1 specimen with the organs of the host; Bismarkburg, Togo; collector and date not recorded; from the mongoose Herpestes sp. (Mammalia: Herpestidae) or Manguste/Mungo ; label bears the inscription No. 27, but is otherwise illegible. ZMB 48072; 2 specimens with the organs of the host; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscription Fundort s. Liste Nr. 26 and Zoologisches Museum Berlin, Jr. No. 1168/10. ZMB 48073; 2 specimens; Langenburg, Tanzania; leg. Fçlleborn, II.1898; host not recorded; label not fully legible and bears the inscription Vom Boy im Magen (?) eines...angeblich gefunden... Langenburg, eingegangen 7.XII.[18]98 [apparently found by a boy in the stomach of a..., received Langenburg 7.XII.[18]98] and No. 12. ZMB 48074; numerous specimens with the organs of the host; Tanga, Tanzania; leg. [Oscar] Neumann, date not recorded; from the bushy-tailed mongoose Bdeogale [crassicauda] puisa Peters, 1852 (Mammalia: Herpestidae) or Buschschwanzmanguste/Dickschwånzige Hundemanguste ; label bears the inscription Bdeogale puisa Ptrs. bei Herpestes, 1 Jahr alt. ZMB 48075; 1 specimen; Cameroon; leg. Schåfer, date not recorded; from the liver of the duiker Cephalophus (Mammalia: Bovidae) or Antilope ; label bears the inscription Fundort: s. Liste No. 13 and Zoologisches Museum Berlin, Jr. No. 1168/10. ZMB 48076; 3 specimens; Bismarkburg, Togo; leg. Conradt, 10.V.[18]93; from the guts of a baboon Papio papio (Desmarest, 1820) [as P. rubescens] (Mammalia: Cercopithecidae); label bears the inscription eing. Ende Mai 94. ZMB 48077; 1 specimen; locality, collector and date not recorded; from the Senegal bushbaby Galago senegalensis Geoffroy, 1796 (Mammalia: Galagidae) or Senegal-Galago/Steppen-Galago ; label bears the inscriptions No. 9, Cavit. Pleural Entozoa and Nr ZMB 48078; 27 specimens; Zoological Garden Berlin; patria host: Tanga, Tanzania, XI.[18]93 ; leg. Oscar Neumann; from the bushytailed mongoose Bdeogale [crassicauda] puisa (Mammalia: Herpestidae) or Buschschwanzmanguste/Dickschwånzige Hundemanguste ; label bears the inscription No. 20. ZMB 48079; 1 specimen with organ pieces; leg. Umlauff, locality and date not recorded; from an antilope pygmaeus [presumably Neotragus pygmaeus (Linnaeus, 1758) (Mammalia: Bovidae)] or Kleinstbæckchen ; label bears the former museum number ZMB 48080; 8 specimens; Cameroon, date not recorded; leg. Schåfer, date not recorded; from the body cavity of a horned viper Cerastes sp. or puff adder Bitis arietans Merrem, 1820 (Serpentes: Viperidae) or Hornviper oder Puffotter ; label bears the inscriptions Fundort s. Liste Nr. 20 and Zoologisches Museum Berlin Jr. No. 1168/10. ZMB 48081; numerous specimens with the organs of the host; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscription Fundort s. Liste unter Nr. 18 and Zoologisches Museum Berlin, Jr. No. 604/1910. ZMB 48082; numerous specimens with the organs of the host; Cameroon, date not recorded; leg. Schåfer, date not recorded; from an unknown ungulate or (Mammalia: Bovidae) or Wiederkåuer ; label bears the inscriptions Zoologisches Museum Berlin, Jr. Nr. 705/11, eing and 1 Milbe lag bei. ZMB 48083; numerous specimens with the organs of the host; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscription Fundort s. Liste unter Nr. 16 and Zoologisches Museum Berlin, Jr. Nr. 604/1910. ZMB 48084; numerous specimens with the lungs of the host; Mombassa, Kenya; leg. Schauer?, date not recorded; from the lungs of an African civet Civettictis civetta (Schreber, 1776) (Mammalia: Viverridae) or Zibetkatze ; label bears the inscription Ein Stçck Lunge mit Wçrmern von einer Zibetkatze gefangen in Mombasa. ZMB 48085; 8 specimens with the organs of the host; Cameroon; leg. Schåfer, date not recorded; from a red river hog Potamochoerus sp. (Mammalia: Suidae) or Pinselohrschwein ; label bears the inscription Fundort s. Liste unter Nr. 23 and Zoologisches Museum Berlin, Jr. Nr. 1168/10. ZMB 48086; 1 specimen; near Kokumiberge, Benue [river], Cameroon; leg. Riggenbach, 27.VII.[19]09; from a baboon Papio sp. (Mammalia: Cercopithecidae) or Pavian ; label bears the inscriptions Zoologisches Museum Berlin Jr. Nr. 1040/09 and in Mue fin... Pavians No. 60. ZMB 48087; numerous specimens with the lungs of the host; Langenburg, Tanzania; leg. Fçlleborn, VI.[18]98; from a python lung Python sp. (Serpentes: Pythonidae) or Python ; label bears the inscription No. 28. museum-zoosyst.evol.wiley-vch.de

14 142 ZMB 48088; numerous specimens with the organs of the host; locality and collector not recorded; 21.II.[19]06; from the mesentery of a prosimian or (Mammalia: Primates) or Halbaffe. ZMB 48089; 5 specimens; Lindi, Tanzania; leg. Fçlleborn, mid February [18]99; from the lung and abdominal cavity of a python Python sp. (Serpentes: Pythonidae) or Python ; label bears the inscription No. 15. ZMB 48090; 5 specimens; Amani, Tanzania [as Deutsch-Ost-Afrika ] leg. Vossler, IV.[19]04; from the lung and abdominal cavity of a puff adder Bitis arietans (Serpentes: Viperidae) or Gemeine Puffotter ; label bears the inscription No. 15. ZMB 48091; 2 specimens; western Africa; collector and date not recorded; from a python Python sp. (Serpentes: Pythonidae) or Python. ZMB 48092; 15 specimens; locality not recorded; Langenburg, Tanzania; leg. Fçlleborn, July; from the lung of a python Python sp. (Serpentes: Pythonidae) or Python ; label bears the inscription No. 4. ZMB 48093; 9 specimens; Lindi, Tanzania; leg. Fçlleborn, mid February [18]97; from the lung and abdominal cavity of a python Python sp. (Serpentes: Pythonidae); label bears the inscription No. 2, date may be incorrect as an almost identical record above comes from ZMB 48094; numerous specimens in liver; Cameroon; leg. Schåfer, 01.VI.[19]11; host not recorded; label bears the inscription J. No. 705/11. ZMB 48095; numerous specimens in intestines and internal organs; Lushoto [as Wilhelmstal ], Tanzania; leg. Erk, date not recorded; from a leopard Panthera pardus (Linnaeus, 1758) (Mammalia: Felidae) or Leopard ; label bears the inscription Pentastomiden im Darm und aus inneren Organen, Sammler: Laboratorium in Daressalam (Dr. Erk) Oberarzt and Zoologisches Museum Berlin, Jr. No. 471/09. ZMB 48096; numerous juveniles; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscription Fundort: s. in Fundortlisten unter Nummer 15 and Zoologisches Museum Berlin, Jr. No. 604/1910. ZMB 48097; numerous juveniles; Cameroon; leg. Schåfer, date and host not recorded; label bears the inscription Fundort: s. in Fundortlisten unter Nummer 17 and Zoologisches Museum Berlin, Jr. No. 604/1910. ZMB 48098; numerous juveniles; Tiko, Cameroon; leg. Piening, 08.IX.1932; from a Fretambo [= a small antilope or duiker] probably Cephalophus sp. (Mamalia: Bovidae); label bears the inscriptions Eing.Nr [entry nr ] and Dr. Panning fragt Kpt. Piening nach Fundort in Wirttier and von einem Fretambo was geschossen in der Messe Lihamba Tier war sehr la... alles besetzt von Wçrmern... Lihamba Kamerun, der 8.IX ZMB 48099; 4 <, 5,; Berlin Aquarium; collector not recorded, 07.III.1939; from a rock python Python sebae (Serpentes: Pythonidae) or Nærdlicher Felsenpython, patria host: Cameroon. ZMB 48100; 31 specimens; Tanzania [as Deutsch-Ost-Afrika ] collector and date not recorded; from a rock python Python sebae (Serpentes: Pythonidae) or Nordlicher Felsenpython ; label bears the inscription No. 25. ZMB 48101; 2 <; Berlin Aquarium; collector, date and host not recorded; label bears the inscription No. 36, Berliner Aquarium. ZMB 48102; 1,; locality, collector, date and host not recorded. ZMB 48103; 1 specimen; locality, collector and date not recorded; from a rock python Python sebae (Serpentes: Pythonidae) or Nærdlicher Felsenpython ; ex Coll. Heymons nr. 36. ZMB 48104; 3 specimens; locality not recorded; leg. Institut fçr vet. Med. Parasitol., date not recorded; from a potto Perodicticus potto (Mçller, 1766) (Mammalia: Lorisidae) or Potto ; Heymons det. ZMB 48105; 1 <, 1,; Cameroon; leg. Schåfer., date not recorded; from the body cavity of a horned viper Cerastes sp. or puff adder Bitis arietans (Serpentes: Viperidae) or Hornviper oder Puffotter ; label bears the inscriptions Fundort: s. Liste No. 20 and Zoologisches Museum Berlin, Jr. No. 1168/10. Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Remarks. The Berlin collection records this species in a wide range of mammalian taxa, including chimpanzees, baboons, bushbabies, pottos, hedgehogs and mongooses. Indeed Riley & Self (1981a) commented that for both A. armillatus and A. moniliformis (see below) almost any mammal can serve as a potential intermediate host. Adults, by contrast, are usually rather hostspecific favouring a few species of large snake. Our data records both vipers and constrictors as definitive hosts. According to Riley & Self (1981a), Armillifer armillatus is restricted biogeographically to Africa, something which is reflected in the collection data noted above. Armillifer armillatus is one of the species known to cause human nymphal pentastomosis in Africa (see e.g. Fain 1966, 1975). Armillifer grandis (Hett, 1915) Material. ZMB 48108; 1,; Democratic Republic of the Congo [as Belgisch-Kongo ]; collector and date not recorded; from the viper Bitis nasicornis (Shaw, 1802) (Serpentes: Viperidae) or Nashornviper ; ex Coll. Heymons nr. 62, label bears the inscription Lula. ZMB 48109; 1 <; Kilo Mines, Ituri [province], Democratic Republic of the Congo [as Belgisch-Kongo ]; collector and date not recorded; from the viper Bitis nasicornis (Serpentes: Viperidae) or Nashornviper ; ex Coll. Heymons nr. 60. ZMB 48110; 2 juveniles; from Uganda; collector and date not recorded; from the viper Bitis sp. (Serpentes: Viperidae) or Puffotter ; label bears the inscription collection Buxton 1a Remarks. According to Riley & Self (1981a), Armillifer grandis is restricted to Africa and this is reflected in the collection data noted above. This is another species known to cause human nymphal pentastomosis (see e.g. Fain 1966, 1975) in Africa. Armillifer mazzai (Sambon, 1922) Figure 4f Material. ZMB 48178, 2 specimens; Papua New Guinea; Ramu-Expedition, date of collection and host not recorded; label bears the inscription No. 11 and Ramu Expedition Eing. 29.I.[18]99. Remarks. This genus Ligamifer Heymons, 1932 was invalidated by Riley & Self (1981a), who transferred its single species, L. mazzai, to the genus Armillifer. According to Riley & Self, the host of the original type material was a brown water snake ; the exact species remaining uncertain. Armillifer moniliformis (Diesing, 1835) Material. ZMB 48112; numerous juveniles; Berlin Zoological Garden; collector not recorded, 18.X.[19]22; from the chest and body cavity of a civet Paradoxurus sp. (Mammalia: Viverridae) or Musang. ZMB 48113; numerous juveniles; locality, collector and date not recorded; from the chest and body cavity of a civet Paradoxurus sp. (Mammalia: Viverridae) or Musang. ZMB 48114; 2 <, 3,; locality and collector not recorded, 08.IX.[19]25; from the carpet python Morelia [as Python] spilota Lac p de, 1804 (Serpentes: Boidae) or Teppichpython ; ex Coll. Heymons nr. 16. museum-zoosyst.evol.wiley-vch.de

15 Zoosyst. Evol. 86 (1) 2010, ZMB 48115; 2 specimens; locality, collector and date not recorded; from a reticulated python Broghammerus [as Python] reticulatus Schneider, 1801 (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 19. ZMB 48116; 1 specimen; Berlin Zoological Garden; collector not recorded, 20.III.[19]34; from an Indian python Python molurus Linnaeus, 1758 (Serpentes: Boidae) or Tigerpython ; ex Coll. Heymons nr. 55. ZMB 48117; numerous juveniles; India; collector and date not recorded; from the civet Paradoxurus sp. (Mammalia: Viverridae) or Musang. ZMB 48118; 1 juvenile; Berlin Zoological Garden; collector not recorded, 04.I.[19]13; from the stomach wall of a loris Stenops gracilis [combination probably invalid] (Mammalia: Lorisidae) or Schlanklori, patria host India; label bears the inscription eing. 20.I.[19]13 [received on 20.I.[19]13]. ZMB 48119; 1 juvenile; Philippines; leg. Jagor, date not recorded; from the Asian palm civet Paradoxurus [hermaphroditus] philippinensis Jourdan, 1837 (Mammalia: Viverridae) or Fleckenmusang ; label bears the inscriptions No. 29 and aus Glas No. 932 (Wçrmer). ZMB 48122; 14 specimens; Java, Indonesia; collector and date not recorded [maybe Damm? see below]; from the small Asian mongoose Herpestes [as Mungos] javanicus (Geoffroy, 1818) (Mammalia: Herpestidae) or Kleiner Mungo ; ex Coll. Heymons nr. 51. ZMB 48123; 6 specimens; Java, Indonesia; leg. Damm, 05.IX. 1923; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 49, label bears the inscription Museum Buitenzorg and mit Longen van Python reticulatus, fix. narme alk. 70 %. ZMB 48124; numerous larvae in host organs; Java, Indonesia; leg. Damm, 16.X.1923; from the liver and stomach wall of the small Asian mongoose Herpestes [as Mungos] javanicus (Mammalia: Herpestidae) or Kleiner Mungo ; ex Coll. Heymons nr. 50, label bears the inscriptions Museum Buitenzorg and fix. alk. 70 % 50 C. ZMB 48127; 1 specimen; India?; collector and date not recorded; probably from an Indian python?python molurus (Serpentes: Boidae) or Tigerpython ; labels poorly legible. ZMB ; 4, 3 and 1 specimen respectively; Berlin Aquarium; collector not recorded, 13.VI.[19]25; from the lung of a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; all labels bear the inscription in der Glasræhre 1 anormales Weibchen (cf. Gesell. Nat. Freunde Berlin 1931) and Zoologisches Museum Berlin, E. K. N ZMB 48131; 3 specimens (1 damaged); Klakah, East Java, Indonesia; leg. H. J. Feuerborn, 15.II.1928; from the lung of a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 13. ZMB 48132, 1 (damaged) specimen; [Berlin?] Aquarium; collector not recorded, 18.VI.[19]31; from the lung of a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython; ex Coll. Heymons nr. 48, label bears the inscription 8 Ju... im Aquarium. ZMB 48133; 5 juveniles; locality, collector, date and host not recoreded [but could be Koch from the Belin Zoo]; label bears the inscription Koch No. 39. ZMB 48134, 1 damaged specimen, Prof. Lange, Reichsgesundheitsamt [Imperial Ministry of Health], VI.1932; encysted in the peritoneum of a Java ape (maybe Macaca fascicularis (Raffles, 1821) (Mammalia: Cercopithecidae)) or Java-Affe ; ex Coll. Heymons nr. 33, label bears the inscription encystiert im Netz eines Java-Affen. Reichsgesundheitsamt Juni 1932 and ded. Dr. Zunker. ZMB 48135, 1 <, locality not recorded; leg. Ruhe-Ahlfeld, 12.V.1924; from the lung of a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; label bears the inscription Zoologisches Museum Berlin, E.K.N ZMB 48136, 3 specimens; Hamburg Zoological Garden; collector not recorded, 14.X.1925; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 20. ZMB 48137; 1,; locality, collector and date not recorded, but presumably Hamburg Zoological Garden (see above); from the lungs of a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 20. Remarks. Again, the final host of this species is a snake; usually a large constrictor. According to Riley & Self (1981a), Armillifer moniliformis is distributed across south-east Asia and this is reflected in the collection data noted above. This is another species known to cause human nymphal pentastomosis (see e.g. Fain 1975), this time affecting populations in the Asian region. Armillifer moniliformis var. heymonsi (Sambon, 1922) Material. ZMB 48120; 9 specimens; Hamburg Zoological Garden; collector not recorded, 30.X.[19]25; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; ex Coll. Heymons nr. 17, label bears the inscription Armillifer moniliformes [sic] var. heymonsi. ZMB 48121; 2 specimens; Berlin Aquarium; collector and date not recorded; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae); ex Coll. Heymons nr. 18, label bears the inscription Armillifer moniliformes [sic] var. heymonsi. ZMB 48125, 1,; locality, collector and date not recorded; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; label bears the inscription Armillifer moniliformes [sic] var. heymonsi. ZMB 48126, 4 specimens; locality, collector and date not recorded; from a reticulated python Broghammerus [as Python] reticulatus (Serpentes: Boidae) or Netzpython ; label bears the inscriptions Armillifer moniliformes var. heymonsi and Nr. 26. Remarks. This taxon, restricted to the reticulated python as its final host (see also data above), was defined rather weakly (e.g. Heymons 1935) on its (typically) smaller females with a pointed posterior end. Its current status is equivocal. In their revision of Armillifer, Riley & Self (1981a: table 1) listed both var. heymonsi and a var. australis as published variants of Armillifer moniliformis. The australis variant was explicitly raised to a separate species by these authors. The status of heymonsi was not addressed further, although Riley & Self cautioned that final measured body size can be influenced by preparation techniques. The variant var. heymonsi may well be a synonym of Armillifer moniliformis, but since we are unable to find a nomenclatural act in the literature formally proposing this, we tentatively list those examples ascribed to heymonsi here separately. Cubirea Kishida, 1928 Remarks. We are not aware of any recent revisions of this genus, nor of data about human infection. Cubirea annulatus (Baird, 1853) Material. ZMB 48051; 1 < juvenile; locality, collector and date not recorded; from the striped sand snake Psammophis sibilans (Linnaeus, museum-zoosyst.evol.wiley-vch.de

16 ) (Serpentes: Colubridae) or Schmuck-Sandrennnatter ; received from Prof. Buxton (Liverpool). ZMB 48052; 1 specimen; Africa; collector, date and host not recorded. ZMB 48053; 1, specimen; Chinguar, Bi Province, Angola; collector and date not recorded; from Anchieta s cobra Naja anchietae Bocage, 1879 (Serpentes: Elapidae) or Anchietas Kobra ; received 1939 from the British Museum London. ZMB 48055; 2 specimens; Africa; collector and date not recorded; from a cobra Naja sp. (Serpentes: Elapidae) or Kobra ; one specimen formerly in the exhibits ( Schausammlung ), one label no longer legible. Cubirea pomeroyi (Woodland, 1921) Figure 4f Material. ZMB 48138; 1 specimen; locality, collector and date not recorded; from the forest cobra Naja melanoleuca Hallowell, 1857 (Serpentes: Elapidae) or Schwarzweiße Kobra ; ex Coll. Heymons nr. 37. Gigliolella Chabaud & Choquet, 1954 Gigliolella brumpti (Giglioli, 1922) Material. ZMB 48106; 1,; Berlin Aquarium; collector not recorded, 25.IV.[19]14; from the Madagascar tree boa Corallus madagascariensis [now Boa mandrita Kluge, 1991] (Serpentes: Aniliidae) or Madagaskar-Hundskopfboa ; ex Coll. Heymons nr. 64. ZMB 48107; 2 juveniles (1 damaged); locality, collector and date not recorded; from the tenrec Centetes [now Tenrec] sp. (Afrosoricida: Tenrecidae) or Tenrek ; ex Coll. Heymons nr. 65. Remarks. Originally in the collection under Armillifer, this species was transferred to their new genus by Chabaud & Choquet (1954). Data from the literature suggests this species is endemic to Madagascar (Gretillat et al. 1962; Slocombe & Budd 1973). Kiricephalus Sambon, 1922 Kiricephalus coarctatus (Diesing, 1850) Material. ZMB 48155; numerous specimens in host organs; Mexico; collector, date and host not recorded. ZMB 48156; 5 specimens; locality, collector and date not recorded; probably from a ratsnake Coluber...? (Serpentes: Colubridae); labels bear inscriptions Wirt: Schlange, Coluber Bemntia, 10135, Pulmo Lamarc Piquo and No. 17. ZMB 48157; 4 specimens; Berlin Zoological Garden; leg. Koch, 03.V.[19]25; from an eastern indigo snake Drymarchon [as Elaphe] couperi Holbrook, 1842 (Serpentes: Colubridae) or Ústliche Indigonatter, patria host North America; ex Coll. Heymons nr. 10. ZMB 48158; 1 specimen; Gorgona, Columbia; collector and date not recorded; from an unspecified snake or Schlange ; ex Coll. Heymons nr. 69, label bears inscription No. 6 and vom British Museum. ZMB 48159; 2 juveniles; locality, collector and date not recorded; from an eastern pine snake Pituophis [as Elaphe] melanoleucus Daudin, 1803 (Serpentes: Colubridae) or Kiefernnatter ; ex Coll. Heymons nr. 38. ZMB ; 3 specimens; Berlin Aquarium; collector not recorded, 1923; from a lung of the eastern indigo snake Drymarchon [as Elaphe] couperi (Serpentes: Colubridae) or Ústliche Indigonatter. ZMB 48162; 4 specimens; Berlin Aquarium; collector not recorded, 31.I.1939; from an eastern indigo snake Drymarchon [as Elaphe] couperi (Serpentes: Colubridae) or Ústliche Indigonatter. Röhlig, D. et al.: Annotated catalogue of tongue worms (Pentastomida) Remarks. A summary of Kiricephalus systematics and biology was published by Riley & Self (1980), with the two species here in the collection regarded as valid and noted as the most common members of the genus. As demonstrated by the Berlin material, K. coarctatus is typically found (as adults) in Neotropical colubrid snakes (Heymons 1935; Montgomery et al. 2006). Kiricephalus pattoni (Stephens, 1908) Figure 5a Material. ZMB 48163; 4 specimens; Berlin Aquarium; collector not recorded, 20.IV.1914; from the Madagascar tree boa Sanzinia [as Corallus] madagascariensis [now Boa mandrita] (Serpentes: Boidae) or Madagaskar-Hundskopfboa ; ex Coll. Heymons nr. 12. ZMB 48164; 1,; Thailand [as Siam ]; leg. Sprater, date not recorded; from the lung of an oriental ratsnake Ptyas [as Zamenis/Coluber] mucosa (Serpentes: Colubridae) or Asiatische Rattenschlange ; ex Coll. Heymons nr. 11. ZMB 48165; 1 specimen; Hong Kong, China; collector and date not recorded; from an unspecified snake or Schlange ; ex Coll. Heymons nr. 71, label bears the inscriptions No. 19 and vom British Museum. ZMB 48166; 3 specimens; from Xiamen [as Amoy ], China; from the lung of an oriental ratsnake Ptyas [as Coluber] mucosa (Serpentes: Colubridae) or Asiatische Rattenschlange ; ex Coll. Heymons nr. 66. Remarks. While Kiricephalus coarctatus is typical for the Americas, K. pattoni is typical for Asia. Heymons (1935) also listed snakes from Madagascar and Australia as hosts and a number of intermediate hosts were identified in India by John & Nadakal (1985). A recent summary of the known definitive (snake) and intermediate (reptilian and amphibian) hosts of K. pattoni was published by Norval et al. (2009). Kiricephalus sp. Material. ZMB 48167; juveniles with entire host; Brasil; leg. Wilh. Ehrhardt, 20.V.1928; from the rodent Hesperomys rufus [current name uncertain]; label bears the inscriptions Hansa Brasil and ZMB 48168; numerous specimens; Berlin Zoological Garden; collector not recorded, III.[19]28; in the mesentery from the gut of a female skunk (Mammalia: Mephitidae) or Stinktier ; label bears the inscription Zoologisches Museum Berlin 976 and im Mesenterium des Darmes. Porocephalus Humboldt, 1811 Porocephalus clavatus (Wyman, 1847) Material. ZMB 48179; 2 specimens (1 undeveloped, 1 clearly ringed); Zoological Garden Hamburg, collector not recorded, 23.X.[19]25; from a northern caiman lizard Dracaena guianensis Daudin, 1802 or Panzerteju/Krokodilteju ; ex Coll. Heymons nr. 41, label bears the inscription es fehlt eine Larve aus Varanus, 1 Porocephalus clavatus irrtçmlich von Vitzthum hinzugetan und von mir weder in Dracaena noch in Varanus gefunden. R. H. [a larva is missing from Varanus, 1 Porocephalus clavatus mistakenly added by Vitzthum but not found by me either in Dracaena or in Varanus R. H.]. museum-zoosyst.evol.wiley-vch.de

17 Zoosyst. Evol. 86 (1) 2010, Figure 5. Further examples of non-type pentastomids in the Berlin collection. a. Kiricephalus pattoni (Stephens, 1908), ZMB 48163, b. Porocephalus crotali (Humboldt, 1808), ZMB 48203, c. Porocephalus stilesi Sambon, 1910, ZMB 48213, d. Waddycephalus teretiusculus (Baird, 1862), ZMB 48230, e. Leiperia cincinnalis Sambon, 1922, ZMB 48169, f. Sebekia oxycephala (Diesing, 1835), ZMB ZMB 48180; 1 specimen; locality, collector and date not recorded, possibly from a caiman lizard Drachena sp. (Squamata: Teiidae) or Krokodilteju. ZMB 48181; 6 juveniles; locality not recorded but presumably the Berlin Zoological Garden (cf. other Koch data); leg. Koch, 15.II.[19].26; from a boa constrictor Boa constrictor Linnaeus, 1758 museum-zoosyst.evol.wiley-vch.de