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Gastrotricha, Chaetonotida) from Australia with the Description of a New Species from Queensland and of Cuticular Ultrastructure in P

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Gastrotricha, Chaetonotida) from Australia with the Description of a New Species from Queensland and of Cuticular Ultrastructure in P Invertebrate Biology 124(2): 119–130. r 2005 American Microscopical Society, Inc. DOI: 10.1111/j.1744-7410.2005.00014.x First record of Polymerurus (Gastrotricha, Chaetonotida) from Australia with the description of a new species from Queensland and of cuticular ultrastructure in P. nodicaudus Rick Hochberga Smithsonian Marine Station, Fort Pierce, Florida 34949, USA Abstract. Identification of specimens in the genus Polymerurus (Gastrotricha, Chaetonotida) is reported for the first time from Australia. The genus includes 3 species from a freshwater lake in southern Queensland. Two species, Polymerurus rhomboides and P. nodicaudus, are well-known cosmopolitans: P. rhomboides is now known from every continent except Ant- arctica, and P. nodicaudus from all continents except Africa and Antarctica. An ultrastruc- tural description of the cuticle in P. nodicaudus is provided and confirms earlier observations on the structure of spined scales in other species of Chaetonotida, suborder Paucitubulatina. The utility of cuticle ultrastructure is discussed in reference to the search for characters de- fining higher-level taxa in the Paucitubulatina. A new species, P. andreae, also cohabits the Queensland lake and is distinguished by the following combination of characters: wide mouth ring; pharynx with terminal bulb; simple spineless scales on the dorsal, lateral, and ventro- lateral body surfaces; and a column of ventral interciliary scales in rectangular, ovoid, and intermediate shapes. Additional key words: gastrotrich, meiofauna, Paucitubulatina, electron microscopy Gastrotrichs are a small but diverse group of mi- only 3 species has been examined despite the vast ar- croscopic worms living in marine, brackish, and ray of cuticular elaborations that exist among species fresh-water habitats, and may constitute an abun- (Rieger & Rieger 1977). dant part of the meiobenthos in both marine inter- The Chaetonotida is traditionally divided into two tidal (Hochberg 1999) and fresh-water lentic systems monophyletic suborders: the Multitubulatina con- (Strayer & Hummon 2001). Schwank (1990) esti- tains a single family and genus, and the Paucitubu- mates B360 species of Gastrotricha within the order latina includes 6 families and 420 genera. The most Chaetonotida, most of which are fresh-water species speciose family of Paucitubulatina is the Chaetono- from lentic habitats, with fewer lotic, marine, and tidae, containing 13 genera and nearly half of all brackish water representatives. Chaetonotidan gas- known species of Gastrotricha. A recent cladistic trotrichs are renowned to be among the smallest analysis identified the Chaetonotidae as potentially known metazoans, generally o1 mm in total body paraphyletic (Hochberg & Litvaitis 2000), while mo- length, and some as small as 80 mm. Despite this min- lecular results using SSU rDNA have hypothesized iscule size, many chaetonotidans are recognized to both polyphyly (Zrzavy 2002) and monophyly (To- have both a complex anatomy and lifecycle (see Rup- daro et al. 2003); the latter molecular studies included pert 1991; Weiss 2001). Their anatomical complexity few genera and species. To date, no convincing mor- is reflected partly in the ornamentation of the cuticle, phological synapomorphies that define the Chaeto- wherein various spines and scales cover much of the notidae have been identified; however, it may be body and may aid in defense and/or flotation. Some reasonable to hypothesize that such characters may of this complexity has been characterized at the ul- be found in the complexity of the cuticle. Still, such trastructural level, but to date the fine structure of synapomorphies are unlikely to be found in cuticle ornamentation because variation in ornamentation appears to be highly pervasive, as noted in a family a Author for correspondence. diagnosis by Kisielewski (1991): ‘‘Body covered, E-mail: [email protected] at least partly, with simple scales, spined scales, 120 Hochberg Invertebrate Biology vol. 124, no. 2, spring 2005 Polymerurus from Australia 121 pedunculated scales or scales bearing lamellae, or, oc- buffer for 24 h, transferred to glycerine on a glass casionally, with a layer of soft and uniform cuticle.’’ slide, sealed with nail polish, and deposited in the In- The genus Polymerurus includes the species with the vertebrate Zoology section of the Queensland Muse- largest described individuals within both Paucitubulat- um. Photographs of live and fixed specimens were ina and Chaetonotidae, ranging 260–770 mm long, and taken with an Olympus BH2 compound microscope is well represented in fresh-water systems worldwide equipped with DIC/Nomarski optics and a Pana- (Kisielewski 1991). Members of this genus are recog- sonic CP610 digital camera. Measurements of gastro- nized by their elongate body and long, superficially trichs were obtained with an ocular micrometer. For segmented posterior furca. The cuticular covering analysis of ultrastructure, gastrotrichs were fixed in of the body is highly diverse, consisting of simple 3% glutaraldehyde in 0.1 M cacodylate buffer (pH flat scales, simple spined scales, stalked scales, and 7.2) for 24 h, followed by a buffer rinse (15 min  4) stalked spined scales, all with various geometric and postfixation in 1% OsO4 in 0.1 M cacodylate shapes. Estimates by Ruppert (1988) and Kisielewski buffer for 1 h. Specimens were rinsed in buffer (1991) place B16–17 species in the genus, most of (15 min  4), dehydrated in a graded ethanol series, which possess spined scales. Polymerurus nodicaudus transferred to propylene oxide, and embedded in an (VOIGT 1901) and P. rhomboides (STOKES 1887) are Araldite/Epon mixture (Procure 812 substituted for the 2 most commonly occurring species in fresh-water Epon; Proscitech, Australia). Resin blocks for trans- systems and are hypothesized to be the most geo- mission electron microscopy (TEM) were sectioned graphically widespread cosmopolitan species. Both at 60 nm with a Diatome diamond knife on a Reic- species have been described from Asia, Europe, and hert or Sorvall ultramicrotome, collected on coated North and South America (e.g., Schwank 1990; grids, and stained with uranyl acetate and lead cit- Kisielewski 1991; Lee & Chang 2000), with P. rhom- rate. Grids were examined with a JEOL 1010 TEM at boides also known from parts of Africa (unpubl. the Centre for Microscopy and Microanalysis, Uni- data). Prior to this study, neither species has been versity of Queensland, and digital photographs were reported from Australia or Antarctica. obtained with a Megaview III color CCD camera and During an investigation of Australian fresh-water processed with analySISs (Soft Imaging System). meiofauna, 3 species of Polymerurus were encoun- Additional grids were examined with a JEOL 100 tered in an artificial lake in southern Queensland, on CX II TEM at the Smithsonian Marine Station at the campus of the University of Queensland in Bris- Fort Pierce, Florida. For light microscopy observa- bane. Based on morphological characteristics, 2 of tions, 1 specimen of P. nodicaudus was sectioned lon- these species are cosmopolitans and the third is as yet gitudinally at 1.5 mm and the sections were stained undescribed. The objectives of the present paper are with Richardson’s stain. Digital photographs were twofold: to provide the first description of Polymer- taken with a Nikon CoolPix 995 on a Zeiss com- urus, including a new species, from Australia and to pound microscope. provide the first ultrastructural account of the cuticle in the genus, with the intention of identifying poten- tial synapomorphies for the Chaetonotidae. Results Methods Taxonomic account Order Chaetonotida REMANE 1924 Specimens of Polymerurus nodicaudus, P. rhom- Suborder Paucitubulatina D’HONDT 1971 boides, and the new species were collected from wa- Family Chaetonotidae GOSSE 1864 ter and vegetation along the shore of a fresh-water Genus Polymerurus REMANE 1927 lake on the campus of the University of Queensland, St. Lucia, Brisbane (B271300S, 153100E). Live gastro- Polymerurus andreae nov. sp. (Figs. 1, 2) trichs were sorted using a dissecting microscope. Voucher specimens for the Queensland Museum Material. Five specimens examined alive as were fixed in 3% glutaraldehyde in 0.1 M cacodylate whole mounts. Two mature whole-mount specimens ’ Fig. 1. Specimens of Polymerurus andreae nov. sp. viewed with light microscopy. A–F are optical sections of the dorsal, medial, and ventral regions of the body. A. Dorsal view of head region. B. Medial view. C. Ventral view showing interciliary, rectangular scales. D. Dorsal view of mid-trunk region showing overlapping simple scales. E. Medial view of pharyngeointestinal region. F. Medial view of trunk region. G. Holotype mounted in glycerine. Cephalion (cp); hypostomion (hy); pleurion (pl); rectangular scale of interciliary region (rs). Invertebrate Biology vol. 124, no. 2, spring 2005 122 Hochberg Fig. 2. Illustrations of features visible in type specimens of Polymerurus andreae nov. sp. A. Ventral view of anterior body region. Scale shapes present on ventral midgut region (mg) are also shown. B. Dorsal head view. Scale shapes present on dorsal body surface are shown. C. Dor- sal view of posterior caudal furca. Cephalion (cp); hypostomion (hy); pleurion (pl); pharyngeal region (ph). prepared as types (QM G218781, QM G219324) and specimen, the caudal furca is 115 mm long. The seg- deposited in the Queensland Museum, South Bris- mented portion of the furca is r90 mm long and bane, Australia. composed of 19–21 segments; the terminal segment Etymology. The species is named in honor of Ms. is the longest, 15 mm. Two columns of ventral loco- Andrea Crowther, Research Assistant at the Queens- motory cilia begin just beyond the hypostomial ridge land Museum, who provided invaluable assistance and extend down the length of the body. with this research. The body is covered by simple spineless scales of Diagnosis. Polymerurus with total body length varying shape (Fig. 1). The dorsal and lateral cutic- 500–710 mm. Head well defined with wide mouth ular covering is composed of flat, rhomboid scales rim and head plates in the form of cephalion, paired with rounded edges (Figs. 1, 2). All scales lack a stalk pleuria, and ventral hypostomion.
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