Seven new species of cestode parasites (Neodermata, Platyhelminthes) from Australian birds

. Cestodes (Platyhelminthes) from Australian birds from the South Australian Museum collections were studied with a focus on common endemic terrestrial hosts. Despite the variable state of preservation of the examined worms, we could identify several new taxa, including Sobolevitaenia whittingtoni sp. nov. and Spiniglans beveridgei sp. nov. (Dilepididae) in Corvus mellori Mathews, 1912 (Corvidae); Notopentorchis musealis sp. nov. (Paruterinidae) in Hirundapus caudacutus (Latham, 1801) (Apodidae); Monopylidium australiense sp. nov. (Dilepididae) in Menura novaehollandiae Latham, 1801 (Menuridae); Dictymetra gerganae sp. nov. (Dilepididae) in Podargus strigoides (Latham, 1802) (Podargidae); Dictymetra longiuncinata sp. nov. in Esacus magnirostris (Vieillot, 1818) (Burhinidae) and Cracticotaenia adelaidae sp. nov. (Metadilepididae) in Gymnorhina tibicen (Latham, 1801) (Artamidae) and Corcorax melanorhamphos (Vieillot, 1817) (Corcoracidae). Several other presumably new taxa that cannot be fully described are also reported. This diversity found in common hosts suggests the presence of a rich, and presently almost completely unknown, fauna of cestode parasites in Australian birds. As field collection permits allowing to explore this fauna are extremely difficult to obtain, this is a demonstration of the usefulness of museum collections to describe at least part of it.


Introduction
With close to 1000 species, including over one third of endemics, the Australian bird fauna is highly diverse and very particular (Lepage 2017).The cestode parasites of this avifauna, especially those of terrestrial endemic species, remain however very poorly known.In their review, Mawson et al. (1986) list about 30 cestode species in these hosts and only a few more have been described more recently (O'Callaghan et al. 2000(O'Callaghan et al. , 2001)).Unfortunately, for administrative reasons, it is presently difficult or even impossible to collect wild birds with the specific objective to survey their parasitological diversity in this country.Thus, the main, or possibly, the single opportunity to study this component of the Australian biodiversity is to access older materials that are conserved in museum collections.Although obviously a second-best option, because of the quality or representativeness of the available material, this approach may allow for some interesting discoveries when old collections have been preserved in good condition.Presently, the single large indigenous Australian parasite collection is maintained in Adelaide in the South Australian Museum (SAMA).It includes historical specimens like those of T.H. Johnston (early 20 th Century) or more recent material like those of I. Beveridge.Due to the reasons mentioned above (and possibly also because of the lack of specialists or of recent interest) almost all of the bird parasites in the Adelaide collections are relatively old and part of them is in poor condition.In the course of a visit organized within the framework of a large international collecting effort (Caira et al. 2017), the senior author reviewed this collection, particularly focusing on parasites from Australian endemic species.This allowed for the identification of a number of new taxa that are described below.The fact that a wide variety of novelties were found in a limited number of hosts is a hint that the Australian cestode fauna in birds is most certainly still largely unknown.

Material and methods
The best available material from hosts of interest was selected during a stay of the senior author at the South Australian Museum, Adelaide, in January 2014.In addition to a small number of stained and mounted specimens, we prepared additional slides from ethanol preserved material using Mayer's hydrochloric carmine staining, dehydration in increasing ethanol series, clearing in eugenol (clove oil) and mounting in Canada balsam.Some scoleces were squashed in Berlese's medium to facilitate examination of rostellar hooks.
All measurements are given in micrometers (if not indicated otherwise) as the range, with the mean and the number (n) of measurements or counts in parentheses.No prevalences are given because these cannot be derived from the available material.Intensities are approximate as most of the material is heavily fragmented.All specimens are deposited in the Australian Helminthological Class Cestoda van Beneden, 1848 Order Cyclophyllidea van Beneden in Braun, 1900 Family Dilepididae Fuhrmann, 1907Genus Sobolevitaenia Spasskaya & Makarenko, 1965 Sobolevitaenia whittingtoni sp.nov.urn:lsid:zoobank.org:act:0B872F99-8193-4EC4-AB17-3A3A0FD00D3F Figs 1-6, Table 1 Etymology The species is dedicated to our colleague and friend, the late Professor Ian Whittington, principal research scientist at the South Australian Museum, Parasites Department.

Paratypes
AUSTRALIA: six specs, same data as for holotype .

Intensity
About 55 specimens in a single host, together with S. beveridgei sp.nov.

Remarks
The presence of spines on suckers, the topology of genital organs in mature proglottides, the pattern of the uterine development and the thick embryophore of eggs unambiguously link our material to the genus Sobolevitaenia Spasskaya & Makarenko, 1965 erected for two species found in Motacillidae in Russia (Spasskaya & Makarenko 1965).This genus comprises about 15 species found in various passeriform birds, mostly Motacillidae and Turdidae, although Corvidae are also known as hosts (Table 1).Spasskaya & Spasskii (1977) listed seven species while Kugi (2000) reported 12 species in the genus.Hosts are all from Northern Hemisphere (Spasskaya & Spasskii 1977;Kugi 2000), except S. korochirei (Voser & Vaucher, 1988) that has been found in Paraguay (Voser & Vaucher 1988).Bona (1994), however, noted the probable cosmopolitan distribution of this genus.
Sobolevitaenia spp.have a homogeneous morphology, including about 20 hooks on two rows for most of them.In order to facilitate comparisons and because metrical reports are highly variable for most species (see e.g., Spasskaya & Spasskii 1977), values from original descriptions are reported as much as possible in Table 1.Most known species show rostellar hooks that are of different size than observed in our material.Furthermore many of them also have a smaller number of testes and (or) cirrus-sacs of different sizes.These characters allow the easy differentiation of our material from the majority of the known Sobolevitaenia spp.(see Table 1).A few remain however more difficult to distinguish:   (Spasskaya, 1958) according to Spasskaya & Spasskii (1977), *** additional measurements in parentheses from Dubinina (1953) in Matevosyan (1963).-Sobolevitaenia iola (Lincicome, 1939), a North American taxon originally reported from Turdus migratorius Linnaeus, 1766 (Turdidae), but also reported from corvids Cyanocitta cristata (Linnaeus, 1758) in Canada (Cooper & Crites 1974).Although resembling our material, S. iola has fewer testes (13-17), slightly shorter hooks (32 long on average) and an armed cirrus (Lincicome 1939).-Sobolevitaenia japonensis Kugi, 2000 from Turdus naumanni Temminck, 1820 in Japan has both a smaller rostellar sac (178-180 × 63-65) and cirrus-sac (96-120 × 1518) as well as spinose cirrus (Kugi 2000).-Sobolevitaenia spinosocapite (Joyeux & Baer, 1955) from various passerines in France, shows a longer cirrus-sac (180-250).Furthermore, Joyeux & Baer (1955) report up to 30 rostellar hooks in this species.Minute spines are mentioned not only on the suckers but on the whole scolex.Joyeux & Baer (1955) also reported this taxon from a corvid, Garrulus glandarius (Linnaeus, 1758), but with noticeably different measurements, including longer (50) and more numerous rostellar hooks (25-28).-Sobolevitaenia unicoronata (Fuhrmann, 1908) from a number of Turdus spp., has hooks of different shape; the length of hooks is similar on both crowns and therefore slightly longer (48) for the second row than those in our material and the double row of hooks is indistinct.Fuhrmann (1908) reported hooks in a single row but Clerc (1911) found them in 2 rows and this point remains contentious.Mettrick's (1958), and numerous other reports of this species in the Palaearctic probably refer to different taxa (Spasskaya & Spasskii 1977).

Species
Two other species initially described from Greenland by Krabbe (1869) possibly belonging to Sobolevitaenia resemble our material, but their exact generic assignations remain unsettled.However, both S. (?) borealis (Krabbe, 1869) and S. (?) trigonocephalum (Krabbe, 1869) have much shorter rostellar hooks and fewer testes.Furthermore S. (?) borealis has an armed cirrus (Baer 1956).We conclude that our material differs from all described species of the genus and we recognise it as a new species, Sobolevitaenia whittingtoni sp.nov.
We should, however, consider that the definition of Sobolevitaenia, based essentially on the presence of spines on the suckers, makes the exact content of this genus uncertain as these structures are easily lost or overlooked.The diversity of the genus is thus probably greater than presently known and, possibly, some taxa currently recognised as members of the genus Monopylidium Fuhrmann, 1899 should be reexamined as possible further species of Sobolevitaenia.

Etymology
The species is dedicated to Professor Ian Beveridge, a friend and colleague, collector of this material, in recognition of his major contribution to systematic helminthology.

Paratypes
AUSTRALIA: 10 specs, same data as for holotype .

Intensity
About 55 specimens in a single host, together with S. whittingtoni sp.nov.

Remarks
The genus Spiniglans Yamaguti, 1959 has been discussed by Bona (1994), Salamatin (1999) and Kornyushin et al. (2009).The latter authors briefly reviewed the generic characters and listed 4 valid species, from Corvidae in the Old World, as well a few other closely related taxa possibly belonging to the genus.They also predicted that more taxa in this group would be found both in other European and tropical corvid species.
Our material can easily be distinguished from most presently recognized species in the genus (Table 2).It differs from S. microsoma (Southwell, 1922) by having more testes; from S. affinis (Krabbe, 1869) by smaller rostellar hooks, fewer testes and a smaller rostellum; from S. constricta (Molin, 1858) by a larger cirrus-sac, rostellar pouch and rostellum; from S. pirinica (Georgiev, Kornyushin & Genov, 1987) by much shorter hooks and a smaller cirrus-sac.Our material most closely resembles S. sharpiloi Kornyushin et al., 2009 but can be distinguished from it by slightly larger rostellar hooks and a longer rostellar pouch and rostellum.It is also similar to S. corvi (Joyeux, Baer & Martin, 1937) but it has more testes and a shorter cirrus-sac.In consequence, we consider it to belong to a new species, Spiniglans beveridgei sp.nov.
Up to now, Spiniglans was known from the Old World only and its host range essentially limited to the Corvidae although the type species was reported in granivore passerines (Emberizidae and Ploceidae) in captivity (Southwell 1922).Its presence in Australia may be due to a recent importation as Corvus colonized (or re-colonized) this continent after diversifying in the Palaearctic (Ericson et al. 2005;Haring et al. 2012).The Corvidae includes around 130 species and is cosmopolitan.It is therefore expected that a larger diversity of parasites of this genus is still unknown, especially in Asia.It remains to be seen whether New World corvids are also hosts of these cestodes, as hinted by the brief report of Pfaffenberger & Butler (1981) of S. constricta ("Anomotania constricta") in Corvus cryptoleucus Couch, 1854 in New Mexico.If confirmed, many more Spiniglans spp.are likely to be discovered in American corvids as the family is highly diversified in the Americas.

Remarks
This material belongs to the genus Neoliga Singh, 1952 as defined by Bona (1994), a genus specific to swifts and tree swifts (Apodidae) and known from Morocco to India, Java and Siberia.Four species are listed by Spasskaya & Spasskii (1977), but the present material does not allow for a detailed comparison.
There is no known species of this genus in Australia, but given the migratory pattern of the whitethroated needletail it is quite possible that parasites from Siberian hosts could be found in Australia.This confirms the prediction of Bona (1994) that the genus distribution is probably wider than presently known.
Paratypes AUSTRALIA: about 24 specs, same datas as for holotype .

Site of infection
Small intestine.

Intensity
About 25 specimens in a single host.
Vitellarium central, oval, mostly regular, composed of few compact lobes.Ovary central, massive, biwinged, multilobate, filling entire width of median field when fully developed, sometimes even slightly crossing lateral osmoregulatory canals.Mehlis' gland central, overlapping seminal receptacle but not always distinct.Seminal receptacle elongated, anterior to vitellarium and between wings of ovary.Vagina thin-walled with discrete basal muscular sphincter; opening posteriorly to cirrus-sac, sometimes partly overlapping it; lined with thin spines in its distal copulatory part (Figs 14-15).
Uterus initially broadly reticulate, developing quickly over whole median field width and extending in lateral field.With further development, uterus becoming compact, saccular, with few deep lobes.Oncospheres slightly oval, 19-24 (21, n = 20) in diameter.Eggs with thin outer shell provided with two big and distinct nuclei; embryophore spherical, thick (Fig. 16).Central embryonic hooks 11 long, lateral embryonic hooks 10 long.

Observations on additional material (not belonging to the type series)
AHC 22931: about 40, extremely small specimens, shorter than 2 mm, maximum width 510, composed of fewer than 20 proglottides but with extremely rapid development.Some early gravid proglottides present.General characters in accordance with the above description to the exception of slightly wider rostella [85-105 (95, n = 10)] and slight variation in number of rostellar hooks (single specimens with 19, 22 or 23 rostellar hooks).
AHC 29271: two slides with fragments of generally poor quality.The observable characters fit the above description.
AHC 29281: a single slide composed of fragments of smaller worms, similar to material from host 22931.Slightly smaller scoleces (200-225 in diameter).

Remarks
In its report on helminth parasite of lyrebirds in Australia, Johnston (1913) described a new acanthocephalan but no cestode.Much later, McOrist & Scott (1989) examined superb lyrebirds suffering from enteritis that was caused by acanthocephalan infections in Southern Victoria and noted the presence of "occasional cestodes of the genus Choanotaenia" in the specimens they studied.To the best of our knowledge, these are the only helminthological reports for these birds.
To our knowledge, there are only two reports of a Monopylidium in Australia.Johnston (1909) mentioned the cosmopolitan species M. passerinum (Fuhrmann, 1907) in Passer domesticus L. in New South Wales and gave a detailed description.This species differed from the present material by several characters including significantly shorter rostellar hooks (Table 3).Monopylidium fieldingi has also been reported from Australia in the butcherbird Cracticus destructor Temminck, 1824[now C. torquatus (Latham, 1801)] in North Queensland (Maplestone & Southwell 1923).However, this taxon has a clearly different anatomy, in particular the presence of 80 rose-thorn shaped hooks in two rows, and is now placed in the genus Cracticotaenia Spasskii, 1966 (Metadilepididae) (see below).
Here, we follow the concept of Spasskii (1993) as updated by Bona (1994) for the definition of Monopylidium.Thus, Icterotaenia Railliet & Henry, 1909 is considered a synonym and some species validated by Spasskii (1993) are now placed in different genera (like Cuculincola Bona, 1994or Spasspasskya Bona, 1994).Globally, about 25 species of Monopylidum are presently recognized (Table 3), although the status of some of them remains equivocal.Our material can easily be differentiated from nearly all of them, although M. attenuatum (Dujardin, 1845) and M. tandani (Singh, 1964) present relatively similar characteristics.However, the former shows a smaller rostellum bearing hooks of different shape, larger oncospheres and embryonic hooks than those of our specimens.It may also have unarmed cirrus (Spasskaya & Spasskii 1977).The latter is a short worm with few (< 20) proglottides, hooks with longer blades and shorter handles and an unarmed cirrus.It is only known from Myophonus caeruleus temminckii Vigors, 1831, a trush found in the Himalayan region (Singh 1964).
Consequently, Monopylidium australiense sp.nov. is the single known cestode parasitizing this endemic host.

Description
This material is in poor state of conservation with contracted and (or) macerated partial strobila and no intact scolex.Scolex width 300-370 (338, n = 3) in diameter; suckers muscular 125-208 (164, n = 12) in diameter.Rostellum 225-255 × 85-137.A single scolex bears an incomplete series of hooks with length 21-25 (23.5, n = 8) (Fig. 39).Worms are much larger and wider than M. australiense (up to 26 mm long and 2.6 mm wide observed, although complete worms should be even larger).The overall anatomy is similar, in particular the few observable rostellar hooks are of the same size and shape as in M. australiense, but, among the observable characters, it seems to have more testes (ca 35-40), a vagina lined with setae and a slightly larger cirrus-sac.The cirrus is armed with well-marked small spines.The available material does not allow for a description but hints at the possible diversity of Dilepididae within lyrebirds.
Paratypes AUSTRALIA: about 14 specs, same data as for holotype .

Site of infection
Intestine.

Intensity
About 15 specimens in a single host.

Description
Body of medium size, with length 65-73 mm and maximum width 1.88-2.3mm (2.03 mm, n = 5) at level of gravid proglottides.Most complete specimens consisting of 79-92 proglottides.Proglottides craspedote, wider than long, progressively becoming longer than wide and up to over twice as long as wide when gravid.Osmoregulatory canals mostly indistinct due to the slight maceration of the specimens.
Testes 60-77 (70, n = 16) in number, in single layer with occasional overlaps, in one continuous large field, filling posterior two thirds of median field, extending in anterior direction up to level of vitellarium both porally and antiporally, often more anteriorly, up to level of ovary on antiporal side (Fig. 19).
Internal vas deferens forming several coils.Cirrus cylindrical, terminally ornate, with long thin setae, mostly difficult to distinguish from those of ductus masculinus.
Vitellarium slightly posterior to level of cirrus-sac, poral to proglottis' midline, antiporal to seminal receptacle, compact and weakly lobate.Ovary small, anterior, slightly poral, bi-alate and lightly lobulate: antiporal wing larger and more anterior than poral wing, at level of vas deferens; poral wing smaller and at level of vitellarium; ovarial wings on both sides of vitellarium.Seminal receptacle elongate, poral to or partially overlapping poral wing of ovary.Vagina opening posteriorly to male pore, most often straight, sometimes slightly bent, rather wide (18-30 in diameter) and short.Vaginal walls regular, bordered externally by loose small cells.

Observations on additional specimens
Although there is no doubt that this material is conspecific with the one described above, we do not include it in the type series due to its poor preservation state.Its observable general characteristics fit the above description with the exception of the occasional presence of well-marked genital papillae.Three hook preparations allowed for more precise observations and measurements of the rostellar armament.We counted 24 hooks on two complete sets, and 20 on an incomplete one.Hooks measuring 48-52 (49.2, n = 23) and 50-56 (52.8, n = 26) in the anterior and posterior crowns, respectively.

Remarks
Helminth parasites of frogmouths are virtually unknown.As for tapeworms, the single reported taxon from these birds is Proparuterina aruensis Fuhrmann, 1911 from the Aru Islands in Podargus papuensis (Fuhrmann 1911;Georgiev & Vaucher 2003).Unidentified cestodes were however more recently reported in P. strigoides in Queensland by Gelis et al. (2011).
Some confusion exists around the systematic organization of dilepidids with cirrus bearing a terminal tuft of long fine spines.According to Bona (1994), they should be distributed in 12 distinct genera and this author provided diagnoses for each of them.By presenting medium-sized strobila, rostellar hooks in two rows, irregularly alternating pores and a reticulate uterus, our material should be placed in Dictymetra Clark, 1952 that has Charadriiformes as 'typical hosts' (Bona 1994).Egg polar processes that are typical of worms with aquatic life-cycles are found in these taxa.It should be noted that other (including terrestrial) birds are also considered by Bona as hosts of members of Dictymetra.They include Passeriformes (for D. belopolskajae Spasskaya & Spasskii, 1973) and Trogoniformes, although the latter are unpublished.This obviously casts some doubts about the monophyly of the genus.
The genus Dictymetra has been erected by Clark (1952) for cestodes found in North American curlews and has been considered very variably by different authors since then, including as a synonym of Krimi Burt, 1944(see Matevosyan 1963) or Anomotaenia Cohn, 1900 (see Bona 1975;Schmidt 1986).In the review of the Dilepididae that we follow here, Bona (1994) gives a detailed diagnosis of the genus, which is significantly different from the original one from Clark (1952).In accordance with Spasskaya & Spasskii (1978), and contrary to Schmidt (1972), he also considers D. radiaspinosa Matevosyan, 1953(syn. D. numenii Clark, 1952, nec Owen 1949) as its type species.
Liga Weinland, 1857, which is found in terrestrial birds, mostly Piciformes, is another genus with characters close to those of Dictymetra but normally with a small strobila and dorsal genital ducts.Several members of this genus, however, have a longer strobila reaching the size of the present material.
For this reason, we compared our material to the presently recognized species of Dictymetra as well as to the "long" Liga spp.(Table 4).
As for Australia, D. adelaidae (Schmidt, 1972) from the spur-winged plover (Lobibyx novaehollandiae Stephens, 1819 now Vanellus miles novaehollandiae) in Queensland was described by Schmidt (1972) as a member of Lapwingia Singh, 1952, which is now considered a synonym of Dictymetra (see Bona 1994).This species does not fit to the diagnosis of Dictymetra and should be place in another genus.Spasskaya & Spasskii (1978) transferred it to the genus Anomotaenia Cohn, 1900.It is in any case very different from the present material for most usual characters, including the length of hooks (shorter) and number of testes (less numerous).
Our material clearly differs from all recognized species in the genus (Table 4) by several characters, the most obvious being the high number of testes, which is only approached but not matched by D. volvulus (von Linstow, 1906).The latter species has however slightly smaller rostellar hooks and a much shorter cirrus-sac.In consequence, we consider our specimens as belonging to a new species that we name Dictymetra gerganae sp.nov.Even though we are confident that this taxon is new, we should notice that the whole taxonomic organization of this group of genera remains quite uncertain due to the scarcity of reliable information for many species.This will probably lead to their complete revision and the definition of new genera, in which our material may need to be placed in the future.

Etymology
From Latin longus (long) and uncinus (hook) in reference to the large rostellar hooks of this species.
Paratypes AUSTRALIA: about 44 specs, same data as for holotype .

Site of infection
Small intestine.

Intensity
About 45 specimens in a single host.
Vitellarium central, compact, irregularly shaped and slightly lobated.Ovary transversely elongated, bi-alate, lobate, antiporal wing larger than poral wing; the latter mostly covered by vas deferens.Mehlis' gland not observed.Empty seminal receptacle round, becoming oval when filled, reaching up to 135 × 68, close to proximal extremity of cirrus sac.Vagina, large, opened posteriorly to male pore, mostly straight and transverse, parallel to cirrus-sac, thin-walled.No sphincters or other ornamentations.

Remarks
This material is partly macerated, with many fragments contracted and overall in an average state of conservation.This explains the lack of data for several characters such as osmoregulatory canals or Mehlis' gland.Our observations, nevertheless, allow for a description that fits the definition of Dictymetra, in particular because of its characteristic scolex structure and tuft of setae in the genital atrium and on the cirrus.The genus Dictymetra is typically found in various families of Charadriiformes, including Burhinidae (Bona 1994) (see remarks above for D. gerganae sp.nov.) and is cosmopolitan (Table 4).
Our material can easily be differentiated from other species in the genus because of its very long hooks and large rostellum.Similar characters are only found in D. laevigata (Rudolphi, 1919), another parasite of thick-knees, and D. paranumenii (Clark, 1952), a parasite of curlews in North America.It differs from both by its smaller number of testes.Additionally, it can be distinguished from the former species by its larger cirrus-sac, and from the latter by its shorter strobila (Table 4).
In addition, the genus Stenovaria Spasskii & Borgarenko, 1973 also resembles our material.It differs from Dictymetra by relatively minor characters related to genital pore alternation, the position of genital ducts in relation to osmoregulatory canals and the absence of armament of long setae on the cirrus and (or) genital atrium.The genus was erected by Spasskii & Borgarenko (1973) and is presently recognized as valid (Bona 1994, Mariaux et al. 2017).It includes 2 species found in thick-knees: Stenovaria facile (Meggitt, 1927) Spasskaya & Spasskii, 1978and S. falsificata (Meggitt, 1927) Spasskii & Borgarenko, 1973 that deserve to be compared to our material.Both differ from it by their much shorter hooks and smaller strobila.In addition, the former species also shows a smaller cirrus-sac and the latter has a larger cirrus-sac and a smaller number of testes.In consequence we consider the present material as a new species that we name Dictymetra longiuncinata sp.nov.

Site of infection
Small intestine.

Description
Despite its similarity with the above described new species, this material probably belongs to another congeneric species.The available mounted specimens do not however allow for a detailed description.
From the single scolex and few fragments available, the hook arrangement is similar to that of D. longiuncinata but hooks are somewhat larger (about 145 long).The rostellar sac (320 × 140) and rostellum (270 × 90) are smaller.It also has a cirrus armed with long setae but has a higher number of testes, i.e., 29-45 (37, n = 12) -and a smaller cirrus-sac, 107-113 × 25-28 (110 × 27, n = 4).It most probably belongs to Dictymetra but additional fresh material is obviously needed to confirm its status.

Site of infection
Small intestine.

Intensity
Two specimens and a few small fragments.Co-infection with Neoliga sp.
Testes large, spherical, 5-8 (6.5, n = 11) in number, in single posterior and dorsal layer, often overlapping vitellarium and posterior parts of ovary, occasionally overlapping longitudinal osmoregulatory canals.External vas deferens convoluted antero-porally, poral to developing uterus surrounded by intensely-stained cells and forming together with them dense mass in anteroporal part of median field.n = 15), consisting of narrower, almost cylindrical, distal part and spherical thick-walled proximal part, often overlapping poral osmoregulatory canals; internal vas deferens coiled in proximal part of cirrus-sac.Cirrus cylindrical,unarmed,[10][11][12]massive,mostly irregular,n = 10), situated adjacent to posterior proglottis margin.Ovary central, transversely elongate, initially slightly bilobate but rapidly becoming more compact and irregular .Mehlis' gland overlapping or just anterior to vitellarium.Seminal receptacle with rounded distal end and tubular and gradually tapering proximal end.Vaginal pore dorsal to pore of cirrus-sac; most of copulatory part of vagina passing posteriorly to cirrus-sac, straight; lumen encircled by thin cellular sleeve; connected with seminal receptacle by thin and short conductive part (Fig. 32).
Young uterus in mature proglottides well marked, with strong walls, developing antero-dorsally to ovary, initially globular (Fig. 32), often with some irregular incomplete septa.With increasing in size, uterus becoming more irregular, with densification surrounding its anterior wall (Fig. 33); with further development, uterus increasing in size; paruterine organ developing mostly anterior to it but also embracing uterus laterally.No fully developed eggs available.

Remarks
The present material unambiguously belongs to Notopentorchis Burt, 1938 as diagnosed by Georgiev & Kornyushin (1994).This genus has recently been thoroughly reviewed and presently comprises eight species, all of them found in swifts from the Old World (Georgiev & Bray 1991;Dimitrova et al. 2017).
Our material is unfortunately in a rather poor state of conservation and limited as neither a scolex with a full set of hooks nor fully developed gravid proglottides are available.It is, however, sufficient to allow examination of most diagnostic characters.These characters were proved to be sufficiently unique to distinguish it from congeners and to allow for a species description.
So far, no species of Notopentorchis has been reported from Australia.The white-throated needletail Hirundapus caudacutus, hosting the worms described here, however occurs in Central and East Asia and winters in Eastern Australia (Lepage 2017), making this range extension logical.The single occurrence of Notopentorchis in this host was reported as N. vesiculigera (Krabbe, 1882) by Oshmarin (1963) in 15 out 29 birds in the Primorye Region (South-Eastern Russia).Besides its European and African distribution, N. vesiculigera clearly differs from our material by its much larger hooks and higher number of testes.We conclude that the present material belongs to an undescribed species that we name Notopentorchis musealis sp.nov.

Intensity
Six specimens.

Description
Body of medium size, with maximum length 35.5 mm and maximum width 1.3-2.12mm (1.7, n = 6) at level of pregravid or early gravid proglottides.Most complete specimen consisting of 240 proglottides.Proglottides craspedote, wider than long, progressively becoming longer, up to slightly longer than wide when gravid.Scolex rounded, not markedly delineated from neck, with diameter 350-475 (407, n = 8), Suckers unarmed,muscular,rounded,n = 35)  cushion-like 128-188 (149, n = 8) in diameter (Fig. 34).Rostellum bearing double crown of regularly arranged hooks, 52-61 (56, n = 8) in number.Hooks triangular, blade terminating with a short, curved tip.Handles and guards bearing well-developed epiphyses, more developed on guard.Epiphyses of hooks in posterior row clearly larger than those of hooks in anterior row .Anterior hooks 23.5-25 (24, n = 4), posterior hooks [21][22]n = 5) wide; proglottization starting shortly behind neck.Genital pores unilateral (a single side-switch observed), opening at border of first third of proglottis length margin.Ventral osmoregulatory canals up to 60 wide, connected posteriorly by anastomosis in each proglottis.Dorsal osmoregulatory canals up to 12 wide.Genital ducts passing between osmoregulatory canals.Genital atrium small.Internal organs closely packed in short and wide proglottides, their shape and relative position clearly visible only in very relaxed or extended proglottides.
Vitellarium central, compact, often heart-shaped or slightly transversely elongated, variable in shape.Ovary fanlike when not contracted, with deep lobes filling most of anterior part of proglottides.Mehlis' gland central and anterior to vitellarium, connected to seminal receptacle by wide and often clearly visible canal that may become almost as wide as seminal receptacle.Seminal receptacle oval to fusiform, remaining rather small, reaching up to 150 × 50, dorsal and between ovary wings or over poral wing, closely to proximal extremity of cirrus-sac.Vagina opens dorsally and slightly posteriorly to male pore, mostly straight, with faint sphincter (Fig. 37).

Remarks
This material is very similar to Cracticotania fieldingi (Maplestone & Southwell, 1923), the only species know in the genus.Cracticotania fieldingi was described from Cracticus destructor (now C. torquatus, Artamidae) in Queensland and attributed to Monopylidum by Maplestone & Southwell (1923).It was later placed in a newly erected monotypic genus by Spasskii (1966) and transferred to the Metadilepididae by Kornyushin & Georgiev (1994).Our material fits the most recent generic diagnosis of these authors with the notable exception that only a very faint vaginal sphincter is visible.Besides this character, it differs from the single known species in the genus by a smaller number of rostellar hooks (52-61 vs about 80), a slightly fewer testes (12-18 vs 16-21) and the differently-shaped and shorter cirrus-sac (globular vs pipette-like and 90-118 vs 130).It should, however, be noted that we examined the type material of C. fieldingi and the measurements provided by Maplestone & Southwell (1923) do not fully correspond to our observations.Although only two slides, mostly with gravid proglottides, are available, we noted a somewhat longer cirrus-sac (up to 180) and fewer testes (12)(13)(14)(15).This, however, does not prevent the unambiguous separation of our material from C. fieldingi and we place it in a new species which we name Cracticotaenia adelaidae sp.nov.
The specimens found in Corcorax Lesson, 1831 fit the above description of Cracticotaenia adelaidae sp.nov. in Gymnorhina tibicen quite well for most metric and meristic characters that can be observed.The main observed differences are in slightly smaller suckers (123-147 vs 155-198) and possibly more rostellar hooks (62-66 vs 52-61), and the faint vaginal sphincter that could not be observed.Keeping in mind that these observations are based on a limited number of incomplete specimens, we consider for now that these small variations (which may be partially exaggerated due to the poor preservation state of our specimens) are likely to be intraspecific.
Although both belonging to the Corvides (see Jønsson et al. 2016), the hosts of C. adelaidae sp.nov.are not closely related and belong to clades that have probably diverged some 20 to 30 Ma ago (Jønsson et al. 2016;Moyle et al. 2016).The Australian magpie (G.tibicen) and the white winged chough (C.melanorhamphos) have however similar diets and habitats (Russell et al. 2017) as well as overlapping ranges (hosts reported in this study were captured a mere 100 km apart).This relative phylogenetic proximity and quite similar ecology probably largely explain the hosting of shared parasites.

Description
This material is very similar to C. adelaidae described above, in most of its morpho-anatomical characters.A clear difference lies however in the shape and size of copulatory organs.While the cirrussac of C. adelaidae is short and globular, it is much more elongated and cylindrical here, always clearly crossing the excretory canals and 70% longer, reaching 155-193 × 30-40 (176 × 36, n = 20).Another discriminating character is the presence of a massive terminal vaginal sphincter, which is absent in C. adelaidae.Hooks are 23-25 (24, n = 9) and 22-24 (22.5, n = 9) long (Fig. 41).The material is however macerated and difficult to interpret, and it is not possible to associate rostellar hook counts and other measurements to specimens for which anatomy is available.The few available specimens from AHC 36526 include some long ones (up to 39.2 mm), including one with a complete scolex (scolex diameter 525, suckers diameter 200-220, about 88 hooks).A vaginal sphincter is present and cirrus-sacs are of a similar shape as C. fieldingi but larger (235-300 × 38-65).
Globally, this material corresponds to the description of C. fieldingi from Cracticus destructor [= C. torquatus] as described by Maplestone & Southwell (1923) and observed by ourselves (see above).It should, however, be noted that C. fieldingi has not only been described in a different host, but also from a locality in Queensland which is some 1500 km north of the New South Wales stations considered here.The available specimens do not allow for a detailed study, but the variability observed in our different samples suggest a possible diversification of Cracticotaenia in the Artamidae.

Discussion
The rich and diversified avian fauna of Australia is undoubtedly hosting a wide variety of cestodes that we safely assume to reach several hundreds of species.Unfortunately, with a few dozens species reported, this area of the country's biodiversity remains both unknown and unexplored.The fact that our study of the SAMA collections allowed to describe seven new taxa from common and abundant host species allows at least two comments.First, museum collections, even older ones, are important tools for the discovery of new taxa, including common but neglected ones.Second, these discoveries only represent a tiny proportion of the real-life diversity, and museum specimens are no substitutes for the exploration of the living world.The diversity, ease of study and access to diverse sources of characters are only guaranteed by fresh collections.In this work, for example, we did not try any sequencing of our samples, because the limited chances of success with these long-preserved specimens, together with the small amount of available material made us deem such attempt too risky.Obviously, a methodical, state of the art collection program would allow for a much easier access to all categories of characters, including molecular, to describe new taxa.

Table 1
(continued on next page).Comparison of Sobolevitaenia whitingtoni sp.nov.with the other species of the genus Sobolevitaenia.Notes: * in the original description, the species is found in a second host species, with slight variations in measurements; ** synonym of S. anthusi

Table 2 .
Comparison of Spiniglans beveridgei sp.nov.with the other species of the genus SpiniglansYamaguti, 1959.

Table 3
(continued on next page).Comparison of Monopylidium australiense sp.nov.with the other species of the generaMonopylidium Fuhrmann,  1899 and Icterotaenia Railliet & Henry, 1909.See text for the accepted generic concept.Note: * 20, 10 and 19 respectively in original reference, a printing error, as confirmed by the length of cirrus-sac on the original illustration.
A few fragmented specimens of this species are found as co-parasites of Cracticotaenia adelaidae sp.nov. in our material from Queensland.Unfortunately, hooks are lost on the single available scolex.Longest fragment 13 mm, maximum width 1.4 mm at level of gravid proglottides.Up to 61 craspedote proglottides, wider than long except for the most gravid ones.Scolex not clearly separated from strobila, 227 in diameter.Suckers rounded 97-115 in diameter.Rostellar pouch very glandular, 167 × 70, extending past posterior margin of suckers.Rostellum muscular, with well-marked terminal pad, 132 × 72.Neck not marked.Genital pores, opening laterally and very anteriorly, at about 15% of lateral proglottis margin, alternating irregularly in short series.Ventral osmoregulatory canals connected posteriorly in each proglottis by transverse anastomosis.Genital ducts passing between osmoregulatory canals.n= 26)in number, in a single layer and one continuous posterior field, extending antero laterally up to level of vitellarium.Cirrus-sac short and weak, 76-92 × 30-42, not reaching poral osmoregulatory canals.Cirrus armed with a small tuft of setae, often lost in our material.Vitellarium central, compact.Ovary anterior transversely elongated.Seminal receptacle rounded, lateral to vitellarium.Vagina posterior to male pore.Uterus initially labyrinthic.No fully gravid proglottides observed.