Progamotaenia festiva (Rudolphi, 1817) Baer, 1927

Beveridge, I. & Shamsi, S., 2009, Revision of the Progamotaenia festiva species complex (Cestoda: Anoplocephalidae) from Australasian marsupials, with the resurrection of P. fellicola (Nybelin, 1917) comb. nov., Zootaxa 1990, pp. 1-29 : 2-11

Progamotaenia festiva (Rudolphi, 1817) Baer, 1927

( Figs 1–11)

Synonyms: Taenia festiva Rudolphi, 1817 ; Moniezia festiva: Blanchard, 1891 ; Hepatotaenia festiva: Nybelin, 1917 ; Cittotaenia festiva: Theiler, 1924 .

Type host: Macropus giganteus Shaw, 1790 (Marsupialia: Macropodidae ).

Other hosts: Macropus dorsalis (Gray) , M. rufus (Desmarest) , M. robustus Gould , M. parryi Bennett , M. antilopinus (Gould) , Onychogalea unguifera (Gould) .

Site in host: bile ducts.

Types: MNB 1888.

Material examined: From Macropus giganteus (eastern grey kangaroo): types; Queensland: 1 specimen, Townsville ( AHC 20875 (S)); 2 specimens, Hervey’s Range, Townsville ( AHC 29409 (S), 7665 (W); 3 specimens, Woodstock ( AHC 29143 (S), 7361(W); 4 specimens, Harvest Home Stn via Charters Towers ( AHC 22287-9 (S)); 1 specimen, Pallamana Stn via Charters Towers ( AHC 22291 (S)); 2 specimens, Clermont ( AHC 29390 (S)); 2 specimens, Emu Park ( AHC 21499 (S)); 1 specimen, Theodore ( AHC 29397 (S)); 1 specimen, Miles ( AHC 29386 (S)); 2 specimens, Chinchilla ( AHC 29410 (S), 9780 (W)); 1 specimen, 40 km N of Charleville ( AHC 29399 (S)); 1 specimen, 66 km E of Moonie ( AHC 29403 (S)); 1 specimen, 30 km E of Bollon ( AHC 29405 (S)); New South Wales: 1 specimen, 50 km N of Moree ( AHC 29385 (S)); 2 specimens, Weilmoringle ( AHC 29401 (S), 34854 (W)); 1 specimen, 85 km N of Bourke ( AHC 29404 (S)); 1 specimen, 50 km N of Bourke ( AHC 29395 (S)); 2 specimens, Warraweena Stn via Bourke ( AHC 28944, 29373 (S)); 1 specimen, 20 km NW of Nyngen ( AHC 29407 (S), 27265 (W)); 1 specimen, Ebor ( AHC 29384 (S), 10417 (W)); 1 specimen, Armidale ( AHC 29411 (S), 9887 (W)); 1 specimen, Coonabarabran ( AHC 29375 (S)); 1 specimen, Nowra ( AHC 29379 (S), 34850 (W)); 1 specimen, Bondo State Forest ( AHC 22092 (S)); Victoria: 1 specimen, Picola North ( AHC 20994 (S)); 1 specimen, Barmah ( AHC 29393 (S)); 1 specimen, Nariel ( AHC 29387 (S)); 1 specimen, Dartmouth ( AHC 20945 (S)); 3 specimens, Mitta Mitta ( AHC 29398 (S), 12057 (W)); 2 specimens, Boho South ( AHC 29377 (S), 34853 (W)); 1 specimen, Dadswell’s Bridge ( AHC 29378) (S)) ; 1 specimen, Pomonal ( AHC 29380 (S)); 4 specimens, Zumsteins ( AHC 20906 (S)); 3 specimens. Mirranatwa ( AHC 29389 (S), 10888 (W)); 3 specimens, Dunkeld ( AHC 29388 (S), 10886 (W)); 1 specimen, Sutton Grange ( AHC 29402 (S), 19775 (W)); 4 specimens, Fraser National Park ( AHC 20855, 21463 (S)); 8 specimens, Eildon ( AHC 29391-2 (S), 9839, 19065 (W)); 6 specimens, Yan Yean ( AHC 20913 (S)); 3 specimens, Healesville ( AHC 29383 (S), 34852 (W)); 1 specimen, Chum Creek ( AHC 29406 (S), 30374 (S)); 1 specimen, Bacchus Marsh ( AHC 20795 (S)); 1 specimen, Brisbane Ranges ( AHC 20856 (S)); 1 specimen, You Yang Range ( AHC 29400 (S), 10166 (W)); 1 specimen, Staughton Vale ( AHC 29412 (S), 10852 (W)); 1 specimen, Avalon ( AHC 29382 (S)); 1 specimen, Lara ( AHC 29381 (S), 34851 (W)); 2 specimens, Marlo ( AHC 20852, 21446 (S)); 1 specimen, Cape Conran ( AHC 29394 (S), 10428 (W)); 1 specimen, Gippsland Lakes ( AHC 29396 (S)); 2 specimens, Bellbird ( AHC 29408 (S), 9711 (W)); 1 specimen, Mt Hickeyard ( AHC 20810 (S)); Tasmania: 1 specimen, Ross ( AHC 29376 (S), 11028 (W)) .

From Macropus dorsalis (black-stripe wallaby): Queensland: 6 specimens, Warrawee Stn via Charters Towers ( SAM 17953, 22285-6 (S)); 5 specimens, Harvest Home Stn via Charters Towers ( SAM 22279-82 (S)) .

From Macropus parryi (whiptail wallaby): Queensland: fragments, Mt Fox ( SAM 29414 (S)); 1 specimen, Mt Surprise ( SAM 7172 (W)); 2 specimens, Townsville ( SAM 29416 (S), 7186 (W)); 1 specimen, Reid River ( SAM 29417 (S), 7164 (W)); 2 specimens, Mt Elliott ( SAM 29418 (S), 7159 (W)); 4 specimens, Inkerman ( SAM 29419 (S), 7178 (W)); 3 specimens, Mingela ( SAM 29421 (S), 7215 (W)); 2 specimens, Yeppoon ( SAM 28957 (S)); fragments, Keppel Sands ( SAM 29415) ; 4 specimens, Banana ( SAM 29420 (S), 19782 (W)); 2 specimens, Thangool ( SAM 29422 (S)); fragments, Eidsvold ( SAM 20069 (S)); New South Wales: 1 specimen, Rivertree ( SAM 20761 (S)) .

From Macropus rufus (red kangaroo): Queensland: 1 specimen, Devoncourt Stn via Cloncurry ( SAM 29433 (S), 34832 (W)) ; 1 specimen, Kiora Stn via Pentland ( SAM 29440 (S), 34831 (W)) ; 1 specimen, 50 km S of Charleville ( SAM 29437 (S)); New South Wales : 1 specimen, 10 km SE of Bourke ( SAM 29423 (S)) ; 1 specimen, 40 km SE of Bourke ( SAM 29438 (S)) ; 1 specimen, 50 km SE of Bourke ( SAM 29436 (S)) ; 1 specimen, Byrock ( SAM 29426(S)) ; 1 specimen, Sturt National Park ( SAM 29439 (S), 34834 (W) ; 1specimen, Narriara Stn via Broken Hill ( SAM 29434 (S), 34833 (W) ; 1 specimen, Thackaringa Stn via Broken Hill ( SAM 29428 (S), 34836 (W)) ; 6 specimens, Menindee ( SAM 20853, 20868 (S)) ; 8 specimens, Kinchega National Park ( SAM 20854 (S)) ; 6 specimens, Warrananga Stn via Wentworth SAM 20759, 20763 (S)); Victoria : 1 specimen, Lake Cullulleraine ( SAM 29432 (S), 19912 (W)); South Australia : 2 specimens, Wilpoorinna Stn via Marree ( SAM 29429 (S), 8138, 34837 (W)) ; 3 specimens, The Twins Stn via Kingoonya ( SAM 29430 (S), 8848 (W)) ; 1 specimen, Wirrealpa Stn via Blinman ( SAM 29431 (S), 8149 (W)) ; 1 specimen, Leigh Creek ( SAM 29441 (S), 8620 (W)) ; 2 specimens, Myrtle Springs Stn via Leigh Creek ( SAM 29435 (S), 8132 (W)) ; 1 specimen, Coondambo Stn via Kingoonya ( SAM 29427 (S), SAM 8852 (W)) ; 2 specimens, Wallerberdina Stn via Pt Augusta ( SAM 29425 (S), SAM 19798 (W)) ; 1 specimen, Melrose ( SAM 21373 (S)); Western Australia : 4 specimens, Yalgoo ( SAM 29424 (S), 34838 (W), BMNH 2008.6 .3.3 (S)) .

From Macropus robustus (wallaroo): Northern Territory: 2 specimens, 40 km N of Katherine ( SAM 29444 (S)); Queensland : 1 specimen, 24 km E of Georgetown ( SAM 29442 (S)) ; 1 specimen, Bluewater Springs ( SAM 28949 (S)) ; 1 specimen, 83 km N of Charters Towers ( SAM 28960 (S)) ; 1 specimen, 30 km N of Charters Towers ( SAM 28955 (S)) ; 1 specimen, Fletcher View Stn via Charters Towers ( SAM 29452 (S), 7412 (W)) ; 1 specimen, Mingela ( SAM 29449 (S), 34842 (W)) ; 1 specimen, Maida Vale Stn via Mingela ( SAM 29453 (S), 7328 (W)) ; 1 specimen, Koonkool Stn via Hughenden ( SAM 29450 (S), 34839 (W)) ; 1 specimen, Woodbine Stn via Prairie ( SAM 29451 (S), 34843 (W)) ; 6 specimens, 70 km W of Cloncurry ( SAM 29455 (S), 34847 (W)) ; 3 specimens, 76 km W of Cloncurry ( SAM 29456 (S), 34836(W)) ; 4 specimens, Devoncourt Stn via Cloncurry ( SAM 29458 (S), 34844-5 (W)) ; 9 specimens, Oban Stn via Mt Isa ( SAM 29457 (S), 11665 (W)) ; 9 specimens, Pallamana Stn via Charters Towers ( SAM 22307 (S)) ; 12 specimens, Warrawee Stn via Charters Towers ( SAM 19755-6 , 22292-22301 (S)) ; 4 specimens, Harvest Home Stn via Charters Towers ( SAM 22302-22305 (S)) ; 3 specimens, Coolagh Stn via Blackall ( SAM 29447 (S), 34840-1 (W)) ; 1 specimen, 7 km E of Barcaldine ( SAM 29454 (S), 34849 (W)) ; 1 specimen, 17 km E of Jericho ( SAM 29448 (S), 34848 (W)) ; 1 specimen, 20 km N of Augathella ( SAM 29443 (S)); New South Wales : 2 specimens, 30 km E of Armidale ( SAM 20793 (S)) ; 1 specimen, Kingstown ( SAM 20791 (S)); South Australia : 2 specimens, Mt Lyndhurst ( SAM 29445 (S)), 8617 (W)) ; 2 specimens, Hawker ( SAM 21392 (S)) ; 4 specimens, Nonning Stn via Pt Augusta ( SAM 22306 (S)); Western Australia : 1 specimen, Wialki ( SAM 20068 (S)) ; 1 specimen, Barrow Island ( SAM 29446 (S), 22958 (W)) ; 2 specimens, Beverley Springs Stn via Derby ( SAM 21491-2 (S)) ; 3 specimens, Yalgoo ( SAM 28959 (S)) ; 4 specimens, Northampton ( SAM 28956 (S)) .

From Macropus antilopinus (antilopine wallaroo): Northern Territory: 2 specimens, 50 km SW of Katherine ( SAM 28958 (S)) ; 2 specimens, 87 km SW of Katherine ( SAM 28954 (S)) ; 4 specimens, 8 km N of Mataranka ( SAM 29460 (S)); Queensland : 1 specimen, Mt Surprise ( SAM 29461 (S), 7172 (W)) ; 8 specimens, Burlington Stn via Mt Surprise ( SAM 29462 (S), 6644 (W)); Western Australia : 4 specimens, Kimberley region ( SAM 21488, 21490 , 21454 (S), 7663 (W)) .

From Onychogalea unguifera (northern nail-tail wallaby): Western Australia: 2 specimens, Kimberley region ( SAM 21440 (S)) .

Description: Based on specimens from M. giganteus . Slender, elongate cestodes, 154–251 (214, n=5) mm long, maximum width up to 8 mm with 437–578 (476, n=5) segments in gravid strobilae. Scolex 0.683–1.11 (0.913) in diameter, prominently lobed; suckers circular, 0.244 –0.325 (0.294) in diameter. Neck prominent. First mature segment 170–205th (190, n=5). Mature segments craspedote, 1.71–2.80 (2.23) wide, 0.163 –0.569 (0.330) long. Genital atrium small, posterior to middle of lateral segment margin; cirrus sac small, elongate, extending to or barely beyond osmoregulatory canals, 0.163 –0.325 (0.266) long, 0.052 –0.065 (0.060) wide; cirrus unarmed; internal seminal vesicle occupies approx. 2/3–3/4 of volume of cirrus sac, 0.078 –0.202 (0.134) long, 0.026 –0.052 (0.034) wide; elongate, spiral, external seminal vesicle, covered with layer of glandular cells, 0.065 –0.195 (0.131) long, 0.033 –0.117 (0.070) wide; vas deferens coils medially, dorsal to uterus; testes arranged in anterior half of segment between osmoregulatory canals; no prominent space between canals and testis fields; testes arranged in 2 dorso-ventral layers, almost invariably in 2 separate groups. Testes 0.026 –0.072 (0.049) in diameter; testis number 71–93 (82) in each segment. Vagina tubiform, lacking glandular investment, opens to genital atrium posterior to cirrus sac, leads to ovoid to subspherical seminal receptacle 0.091 –0.247 (0.166) x 0.085 –0.182 (0.127). Ovary small, flabelliform, medial to seminal receptacle 0.052 –0.098 (0.072) x 0.033 –0.078 (0.055); vitellarium reniform, posterior to ovary, 0.072 –0.143 (0.102) x 0.052 –0.117 (0.067); Mehlis’ gland anterior and medial to vitellarium. Uteri paired in each segment, transverse, tubular; early uterus extends from beyond level of ovary to near proximal pole of cirrus sac; developing uterus extends medially and laterally, lateral extensions crossing osmoregulatory canals dorsally; medially uteri of each segment do not meet in mid-line. Gravid segments 0.244 –0.683 (0.406) long, 2.93–5.82 (3.88) wide. Egg spherical, shell smooth, 0.045 –0.050 (0.048) in diameter; pyriform apparatus terminating in numerous reflexed filaments; oncosphere 0.013 –0.018 (0.015) in diameter. Osmoregulatory canals paired; ventral canal generally wider than dorsal, medial to it; ventral canal 0.020 –0.085 (0.044) in diameter, dorsal canal 0.010 –0.033 (0.018) in diameter; transverse canal connects ventral canal at posterior margin of each segment; accessory canals present in some specimens.

Remarks. The redescription of this species is based exclusively upon specimens from Macropus giganteus , identified by Rudolphi (1819) as the host of the type specimens. The type specimens were collected from a grey kangaroo, M. giganteus , which died in the Schönbrunn Zoological Gardens, Vienna, and were collected by Bremser and sent to Rudolphi in Berlin. Putting aside the reliability of the identification of species of Macropus in European zoos during the early nineteenth century, the genetic study of Beveridge et al. (2007), based on the cox 1 gene, indicated that specimens of bile duct cestodes collected from this host species over a broad geographical range in eastern Australia belonged to the same genetic clade and therefore potentially to the same species. A wide range of specimens was also available from this host species covering most of its geographical range ( Fig. 12) and all were relatively uniform morphologically. The cestodes were long (> 150 mm), composed of> 400 segments, with the first mature segment being about the 170–205th segment from the scolex. The scolex was usually prominently four-lobed ( Fig.3), though in incompletely relaxed specimens, the lobes of the scolex were shortened ( Fig.2). In the mature segments, the testes were almost invariably separated into two equal groups, with 71–93 testes per segment, and the testis fields abutted the longitudinal osmoregulatory canals. The cirrus sac was small and reached the osmoregulatory canals but rarely extended significantly into the medulla ( Fig. 8). The external seminal vesicle was spirally coiled and was surrounded by gland cells; when filled, several coils were expanded, giving the appearance of a multichambered external seminal vesicle ( Fig. 5).

Specimens of P. festiva from M. rufus formed a distinctive clade in the genetic analysis of Beveridge et al. (2007). Morphologically, these specimens (with a single exception dealt with below) were also uniform throughout their geographic range ( Fig. 14), and were similar to the specimens from M. giganteus , differing only in the more pronounced extension of the cirrus sac into the medulla in some specimens ( Fig. 9). However, there was substantial variation in this feature within specimens from M. rufus and there was no difference in the mean size of the cirrus sac between specimens from M. giganteus and M. rufus (Table 1). Consequently, no significant and consistent morphological differences were observable between specimens from these two host species. Collections from M. rufus were relatively extensive, but were concentrated in the eastern part of the host range ( Fig. 14). Additional collecting is required in central and western Australia to ensure that the genetic clade is uniform across its entire range.

In the genetic analysis, all specimens from M. parryi also formed a distinctive clade. Morphologically, these specimens, which were obtained from across a wide geographical range ( Fig.13), were relatively uniform morphologically, but differed in one feature. In almost all specimens, the external seminal vesicle was fusiform ( Fig. 6) rather than spirally coiled. The testes were in separate groups, a feature of all specimens in this group, except for two specimens, one of which was immature (SAM 29414) and the other was severely contracted (SAM 20069). In all well-preserved specimens examined, there was a distinctive space between the osmoregulatory canals and the lateral extent of the testis fields ( Fig. 10), thereby distinguishing these specimens from those found in M. giganteus and M. rufus .

Specimens from M. robustus and M. antilopinus occurred in four distinct clades in the genetic analysis of Beveridge et al. (2007), and these differences were reflected in part in their morphological features. Specimens collected from M. robustus in South Australia and Western Australia (type 1 in Fig. 15) were morphologically identical with those from M. giganteus and M. rufus in lacking a space between the testis field and the osmoregulatory canals. Specimens from M. robustus from north western Queensland (Mt Isa, Cloncurry) (type 2 in Fig. 15) were distinguishable in having a distinct space between the osmoregulatory canals and the testis fields as in the specimens from M. parryi ( Fig. 10), but were distinguishable from specimens in M. parryi in having a spiral external seminal vesicle. This clade was also found in M. dorsalis in north-eastern Queensland and in M. rufus from Cloncurry and Pentland, Queensland, but in each case, this genetic form was identifiable by the space between the testes and the osmoregulatory canals and by the spiral external seminal vesicle. Specimens with similar morphological features were seen in M. robustus from Armidale, NSW, Barrow Island and the Kimberley, WA, but since no material was available from these locations for genetic analysis, the relationships of these cestodes remain to be delineated. The genetic clade (3 in Fig. 15) identified from coastal north-eastern Queensland was characterised morphologically by the lack of a space between the testis fields and the osmoregulatory canals and was therefore not distinguishable from the clade found in south and western Australia (1 in Fig. 15).

All material examined from M. antilopinus in the Northern Territory and all but one specimen examined from Queensland was morphologically distinctive in having a distinct space between the osmoregulatory canals and the testis fields and a greater number of testes per segment than in other specimens ( Fig. 11) (Table 1). However, the space between testis fields and osmoregulatory canals was somewhat variable and was small in one specimen. This morphological form was also seen in a single M. robustus from the Northern Territory, in an area in which M. antilopinus and M. robustus are sympatric (4 in Fig. 15). No entire specimens of this morphotype were collected and therefore although it appears to represent a distinctive species, it is not described as a new species herein. In the Mt Surprise region of northern Queensland, M. antilopinus and M. robustus are also sympatric ( Fig. 15). The fixed material from M.antilopinus from this area is similar to that from the same host species in the Northern Territory. However, the single specimen of P. festiva from M. antilopinus from north Queensland used in the genetic analysis appears to be identical with the morphological form predominant in M. robustus in that region suggesting that both morphotypes may occur in the two host species in this region. More extensive collections are needed to resolve the morphotypes present in wallaroos in this region.

Metric data of P. festiva from various host species are presented in Table 1. Whilst there are some differences in measurements, particularly for specimens designated as clade 4, initial inspection of the data suggested that detailed statistical analysis was not warranted as it was unlikely to yield useful taxonomic information.

Fragmentary material was available from Onychogalea unguifera from the Kimberley region of Western Australia. While generally referable to P. festiva , additional specimens are required to determine if it represents an additional species.