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Studies on the Systematics and Biogeography of Terrestrial Flatworms (Platyhelminthes: Tricladida: Terricola) of the Australian Region

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Studies on the Systematics and Biogeography of Terrestrial Flatworms (Platyhelminthes: Tricladida: Terricola) of the Australian Region ResearchOnline@JCU This file is part of the following reference: Winsor, Leigh (2003) Studies on the systematics and biogeography of terrestrial flatworms (Platyhelminthes: Tricladida: Terricola) of the Australian region. PhD thesis, James Cook University. Access to this file is available from: http://eprints.jcu.edu.au/24134/ The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owner of any third party copyright material included in this document. If you believe that this is not the case, please contact [email protected] and quote http://eprints.jcu.edu.au/24134/ Chapter 5 Systematic Account and Phylogeny ...I don't care much about working with land planarians. There are just too many of them and the older descriptions based primarily on colour are very difficult to apply to pickled specimens. Libbie Hyman in litt to Carl Pantin, early 1950 (letter in the Pantin Collection, Zoology Department, University of Cambridge, access courtesy of Dr Janet Moore) It is curious that so few Australian zoologists have concerned themselves with the study of the Land-Planarians. This is the more to be regretted inasmuch as the opportunities for collecting these animals are rapidly passing away with the clearing of the bush. Moreover, much remains to be done in the investigation of these and other Cryptozoic animals. The Land-Planarians, in particular, still demand thorough comparative anatomical investigation with a view to revising the generic classification. Dendy (1915) 5.1 INTRODUCTION TO THE FAMILIES, SUBFAMILIES & GENERA The classification of the Terricola provided in the Land Planarian Indices of the World (Ogren, Kawakatsu and others), together with the provisional classification of the Australian Geoplanidae (Winsor 1991c), are taken as the starting points for the taxonomy and systematics of the Terricola of the Australian region in this thesis. A summary of the families, subfamilies, and genera of the Terricola worldwide is provided elsewhere (Table 1.4, Chapter 1) Australia and New Zealand and their subantarctic islands share Terricola taxa at the levels of sub-family and genus, especially Artioposthia sensu lato and Caenoplana. They also share some species through the inadvertent introduction of flatworms such as Parakontikia ventrolineata. For these reasons, and for comparative anatomical purposes, New Zealand taxa are included in this study and accordingly Arthurdendyus, Coleocephalus and Fyfea carnleyi have been revised. The collective group genus Australopacifica Ogren & Kawakatsu, 1991 is not considered here as it is a repository for caenoplaninids about which insufficient internal details are known to permit reliable assignment of these species elsewhere. In order to provide a complete account, all genera present in the Australian region are included here (Table 5.1). Current diagnoses from the literature, emended only to harmonize the terminology 145 used throughout this thesis, are provided for those genera that have not been considered in detail in this study. The following descriptions are accompanied by simple line drawings of a representative species, eye pattern, simple diagram of the copulatory organs, and a map showing the distribution of the genus (solid dots or lines) together sometimes with an indication of tracks connecting them (dashed lines). Detailed descriptions and anatomical illustrations of new and revised taxa are provided in Appendix 8. In order to facilitate later phylogenetic analysis and comparisons between taxa, the format of descriptions adopted here accords with that also used by the team of Dr Ana Maria Leal-Zanchet, Instituto de Pesquisas de Planarias, UNISINOS, Sao Leopoldo, RS, Brazil. The descriptions consider external features, and internal anatomy of the head, pre-pharyngeal region, and reproductive organs. Abbreviations used in figures follow the glossary. 5.1.1 Nomenclature The new taxa described in this thesis are nomina nuda as the provisions of Article 11 of the International Code of Zoological Nomenclature (ICZN 1999) are not met. It is intended to publish the descriptions of new taxa as soon as possible. 5.1.2 Etymology The names of many genera in the Terricola have the suffix —plana, gender feminine, from the Latin planus, flat. The early workers, for example Diesing (1850), did not provide an explanation of the etymology of names in which this suffix was used. It possibly alluded to the common name, planarian, by which free-living triclad flatworms are generally known. It is in this context in which the suffix -plana is used for new taxa in this thesis. 5.1.3 Field Keys to the Terricola in Australia Pictorial field keys, based upon external characters, are provided in Appendix 9, Keys A9.1— 9.5 for the families and genera of Terricola in Australia. 146 Table 5.1. List of Families and Genera of the Australian Region (*refer also to species description in Appendix 8) FAMILY: Rhynchodemidae Subfamily: Rhynchodeminae Cotyloplana Digonopyla Dolichoplana Platydemus Rhynchodemus FAMILY: Geoplanidae Subfamily: Anzoplaninae sub.fam.nov. Anzoplana gen.nov* Fyfea gen.nov* Subfamily: Caenoplaninae Apogaioplana gen.nov.* Arthurdendyus * Artioposthia Australoplana* Barringtoplana gen.nov.* Bassioplana gen.nov.* Caenoplana Coleocephalus * Danidoplana gen.nov.* Fletchamia Kontikia Kynagoplana gen.nov.* Lenkunya* Nelipoplana (Nelipoplana) gen. et sub.gen.nov.* Nelipoplana (Peokrypta) gen. et sub.gen.nov.* Newzealandia Parakontikia Reomkago Sokoplana gen.nov.* Tasmanoplana Thosteelia gen.nov.* FAMILY: Bipaliidae Bipalium 147 SYSTEMATIC ACCOUNT The present three families of the Terricola are insecurely based (Cannon 1986). None of the diagnoses for the families and subfamilies includes a comparable suite of characters at each level. Revision of these diagnoses is outside the scope of this thesis. In my view inclusion of general characters concerning the presence and distribution of eyes are best included in the diagnoses for family, with detailed descriptors used in the diagnoses for genera. Eye distribution patterns provide useful diagnostic characters at the genus level. At the subfamily level the descriptors are broad and serve little diagnostic purpose; the family diagnoses adequately cover the subfamilies. The character for eye type, for which there is unfortunately a paucity of data, is likely to be important at the family or subfamily levels. 5.2. RHYNCHODEMIDAE Diagnosis (Ogren & Kawakatsul988a): Land planarians of elongate cylindroid form with two eyes near the simple, tapered anterior end; no tentacles or headplate; with well-defined creeping sole occupying part of the ventral surface. Anterior end may have a sucker organ on ventral surface (Cotyloplana). 5.2.1 Rhynchodeminae Diagnosis (Ogren & Kawakatsu, 1988a): Rhynchodemidae with strong cortical musculature in which the subepithelial longitudinal muscle fibres are grouped into large, definite bundles; penis papilla absent or greatly reduced (Platydemus in part). 5.2.1.1 Genus Cotyloplana (Plate 5.1) Cotyloplana Spencer, 1892 Cotyloplana Spencer, 1892; Graff, 1899; Ogren & Kawakatsu, 1988a; Winsor, 1997 Diagnosis (Ogren & Kawakatsu 1988a): Rhynchodeminae, flattened body, with wide creeping sole; one sucker, ventrally located, present at anterior of body in front of the creeping sole; copulatory organs similar to Platydemus with large, 148 Plate 5.1. The habitus, eye pattern, copulatory organs, and occurrence of Cotyloplana (Rhynchodemidae: Rhynchodeminae) CZ f g pd pp spacious male antrum having a small, obtuse, penis papilla at its base through which the ejaculatory duct empties; female organ simple, with narrow curved female genital duct, dorsally situated, reduced anteriorly where it opens into the dorsal part of the common antrum; viscid gland (posterior diverticulum) present. Diagnosis (emended, this thesis): Rhynchodeminae with small robust body, flattened, mouth about mid body, with gonopore closer to posterior end than to mouth. Broad creeping sole 60-70% of body width. Anteroventral sucker present, single cup- type, in front of creeping sole. Eyes, large, paired, retina-type. Cutaneous musculature tripartite, with circular, decussate helical, and longitudinal muscles. Longitudinal muscles in strong bundles. CMI 11-14%. Insunk ventrally (in part the sucker retractor muscles). Parenchymal longitudinal musculature strong, concentrated in ventral plate. PMI 13-25%. Anteriorly very strong dorsal longitudinal parenchymal muscles. Supraintestinal transverse muscles, subneural transverse muscles, dorsoventral muscles strong. Pharynx bell-form. Pharyngeal musculature with inner epithelium, longitudinal, mixed circular-longitudinal, then longitudinal muscles; outer musculature with epithelium, longitudinal, circular, then longitudinal muscles; pharyngeal pouch diverticulum may be present. Testes ventral, lateral to nerve cords, uniserial, post-pharyngeal, extend from just anterior to mouth to just before copulatory organs; vasa deferentia ventral, rise, recurve anteriorly, separately enter penis bulb, unite in anterodorsal bulb, open mid anterior bulb into seminal, then prostatic vesicles, with muscular ejaculatory duct. Penis papilla poorly differentiated. Penis eversible-type. Penis bulb strongly muscular, with inner longitudinal, outer circular muscles. Male antrum large, capacious, with distal mucosal folds. Ovaries anterior; ovovitelline ducts unite, form common ovovitelline duct which enters female genital duct posteriorly. Female genital duct short, dorsally
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