Platyhelminthes: Tricladida: Geoplanidae) from Stewart Island, New Zealand
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Tuhinga 20: 23–32 Copyright © Museum of New Zealand Te Papa Tongarewa (2009) A new subfamily, new genus and new species of terrestrial flatworm (Platyhelminthes: Tricladida: Geoplanidae) from Stewart Island, New Zealand Leigh Winsor School of Marine and Tropical Biology, c/- People and Equity Directorate, James Cook University, Townsville, Queensland 4811, Australia ([email protected]) ABSTRACT: Within the context of ongoing taxonomic revisions of terrestrial flatworms from the Australia–New Zealand region: the new genus Eudoxiatopoplana is erected; the new species Eudoxiatopoplana bilaticlavia is described from Stewart Island; and the new subfamily Eudoxiatopoplaninae is erected within the family Geoplanidae. KEYWORDS: Taxonomy, Eudoxiatopoplaninae, new subfamily, Eudoxiatopoplana bilaticlavia, new genus, new species, resorptive bursa. Introduction the stripe pattern follows the scheme of Graff (1899: 25). Determination of the cutaneous and parenchymal muscular To date, only three species of terrestrial flatworms are known indices (CMI, PMI) follows Winsor (1983). Nomenclature from Stewart Island, New Zealand, all of which were found and type of adenodactyl follows Winsor (1998b). The speci - in modified habitat: Arthurdendyus australis (Dendy, 1894), men is held in the collections of the Museum of New from nurseries and agricultural land; and A. testaceus (Hutton, Zealand Te Papa Tongarewa, Wellington (MONZ). 1880) and A. triangulatus (Dendy, 1894), from nurseries (Johns & Boag 2003). In the Museum of New Zealand Te Abbreviations used in figures Papa Tongarewa collection of terrestrial flatworms from the ad adenodactyl Subantarctic and other islands presently being studied by bd bursal duct the author (Winsor 2006), there is a single specimen from cm cutaneous musculature Stewart Island. This paper concerns the description of this cod common ovovitelline duct specimen, considered here to be an endemic new genus and cs creeping sole new species of terrestrial flatworm accommodated in a new dsv distal seminal vesicle subfamily of the Geoplanidae. ed ejaculatory duct fa female antrum fg female genital canal Materials and methods gp gonopore The specimen selected for histology was processed to paraffin ip inverted penis wax, longitudinally sagittally (LSS) and transversely sec - in intestine tioned (TS) at 8μm, and stained using the MSB (Martius ma male antrum scarlet blue) method (Winsor 1998a,b). Nomenclature of mo mouth 24 Tuhinga, Number 20 (2009) nc nerve cord terrestrial flatworms for over 55 years and for her friendship ovd ovovitelline duct with the author for the past 35 years. It also alludes to generic ph pharynx characters – particularly the dorsal testes, eye pattern and php pharyngeal pouch musculature that are usually associated with taxa from the pm parenchymal musculature (ring zone) Neotropical and Oriental zoogeographic regions – that, psv proximal seminal vesicle until now, had not been reported from the Australia–New rb resorptive bursa Zealand region. Suggested vernacular name: Eudoxia’s rh rhammites strange flatworm. sd sperm duct DISTRIBUTION: Stewart Island, New Zealand. sdv spermiducal vesicle sg shell glands Systematic discussion te testis Until recently, the suborder Tricladida (Platyhelminthes) vd vas deferens was divided taxonomically into four ecologically based infraorders: Maricola (marine planarians); Paludicola (fresh - Systematics water planarians); Cavernicola (cave-dwelling planari ans); and the Terricola (terrestrial planarians). However, molecular Family Geoplanidae Stimpson, 1857 analyses of planarian flatworms have caused rethinking of Eudoxiatopoplana new genus these infraorders; whilst the Maricola are clearly mono - DIAGNOSIS: Geoplanid with a small, robust body, lanceo- phyletic, it has been demonstrated that the Paludicola are late, broader posteriorly than anteriorly, sub-cylindrical in paraphyletic, the Terricola being the sister group to one of cross section. Mouth situated just behind mid-body, gono- the three paludicolan families, the Dugesiidae (Álvarez- pore closer to mouth than to posterior end. Eyes in multiple Presas et al. 2008). As a consequence of these analyses a rows around anterior tip and dorso-anteriorly, extending the new higher classification of the paludicolan and terricolan length of the body. Sensorial margin passes antero-ventrally flatworms is urgently required, and will be proposed in the around the anterior tip. Sensory papillae absent. Anterior near future (Ronald Sluys, pers. comm. 2008). Pending the ad hesive cup, sucker absent. Creeping sole non-ciliated, 31% publication of a new classification of planarian flatworms, of body width. Cutaneous musculature tripartite, with cir- the following systematic discussion reflects the current cular, helical and longitudinal muscles; longitudinal muscles traditional taxonomy of the Terricola. in well-defined bundles; dorsal cutaneous musculature almost The Geoplanidae, to which all native terrestrial flatworms double of that ventrally, CMI 4.4–6.5%. Parenchymal lon- of New Zealand and its Subantarctic Islands belong, is gi tudinal muscles present as a massive ring zone, PMI 38.5%, defined as ‘Terricola with numerous small eyes concentrated stronger ventrally than dorsally and intersected by a closely around the anterior, and along the sides, sometimes located woven meshwork of oblique-transverse and dorso-ventral dorsally, or without eyes; without auricular or tentacular muscles fibres. Testes dorsal, extending from behind ovaries organs, or semilunar headplate’ (Ogren & Kawakatsu 1990). to posterior end. Efferent ducts ventral. Penis inverted type. This family presently comprises four sub-families: the Ovaries in anterior body third. Ovovitelline ducts unite then Geoplaninae (Ogren & Kawakatsu 1990: 83), which occur enter female genital canal ventro-posteriorly (postflex con- largely in South and Central America; the Caenoplaninae dition). Female genital canal enters antrum horizontally. (Ogren & Kawakatsu 1991: 28, emended by Winsor 2006), Resorptive bursa present. Adenodactyl present. Viscid gland centred mainly in the Australo-Pacific region; the Pelmato - absent. planinae (Ogren & Kawakatsu 1991: 33) in the Southeast TYPE SPECIES: Eudoxiatopoplana bilaticlavia new species. Asia–Indo-Pacific region; and the Anzoplaninae (Winsor ETYMOLOGY: The name Eudoxiatopoplana is composed of 2006) from Australia, New Zealand and the Subantarctic Eudoxia (Greek = honoured), atopos (Greek = out of place, Islands. strange) and plana (Latin = flat, planarian). Gender: femi- With multiple eyes concentrated around the anterior nine. It honours Emeritus Professor Eudoxia Maria Froehlich end, dorsally and along the sides, and without auricular or (Departamento de Zoologia, Universidade de São Paulo, tentacular organs or a semilunar headplate, Eudoxiatopoplana Brazil) for her outstanding contribution to the study of clearly belongs in the Geoplanidae. However, assignment A new subfamily, new genus and new species of terrestrial flatworm (Platyhelminthes: Tricladida: Geoplanidae) 25 of Eudoxiatopoplana to one of the four subfamilies in the and sperm are digested intraluminally. Four types of resorp- Geoplanidae is problematic as the new genus has a mix of tive bursae are now identified: the posterior-medial resorptive mutually exclusive taxonomic characters. bursa present in Eudoxiatopoplana; and three types of lateral The eye pattern in Eudoxiatopoplana, with lateral cluster - bursae present in other taxa. ing and eyes extending the length of the body, is similar to The three types of lateral bursae are all only lightly that found in some taxa of the Caenoplaninae, Anzoplaninae muscularised: atrial lateral bursae embedded in the atrial and Geoplaninae, but differs from these in that multiple musculature, the short bursal stalks of which arise at the rows of eyes, instead of a single row of eyes, pass around the entrance of the female genital canal in Australopacifica willeyi anterior tip. In the Pelmatoplaninae, the eyes do not extend (Busson, 1903) (Caenoplaninae), which has anterior-facing, around the anterior tip, are crowded antero-laterally and inequi-bilobed, bilateral bursae; parapharyngeal lateral rarely extend beyond the mouth. The eye pattern of the bursae situated on either side of the pharynx, where the anterior end in Eudoxiatopoplana is not dissimilar to that short bursal stalks arise from the ovovitelline ducts, present observed in species in the Bipaliidae, though an expanded in Anzoplana trilineata Winsor, 2006 (Anzoplaninae); and headplate is absent in Eudoxiatopoplana. ovarian lateral bursae, previously termed ‘parovarian tissue’ The presence of dorsal testes excludes Eudoxiatopoplana and at that time of unknown function (Fyfe 1937, 1953), from the Caenoplaninae and Pelmatoplaninae, which have situated at the proximal end of the oviducts and ventro- ventral testes, and from the Anzoplaninae, which have dorso- lateral to the ovaries, now recognised as resorptive bursae ventral testes. The presence of dorsal testes, regarded as an (Winsor & Stevens 2005). These bursae are present in the apomorphy, characterises the Geoplaninae whose distribu - caenoplanines Arthurdendyus triangulatus (Dendy, 1895), tion is confined to the Neotropical zoogeographic region. A. vegrandis Winsor & Stevens, 2005, and the subantarctic The presence of this character state in a species from the taxa Kontikia marrineri (Dendy, 1911), Kontikia ashleyi Australian region is unique. However, Eudoxiatopoplana is (Fyfe, 1953) and Fyfea carnleyi (Fyfe, 1953).