The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894 (Crustacea, Syncarida, Anaspididae)
Total Page:16
File Type:pdf, Size:1020Kb
© The Author, 2016. Journal compilation © Australian Museum, Sydney, 2016 Records of the Australian Museum (2016) Vol. 68, issue number 7, pp. 313–364. ISSN 0067-1975 (print), ISSN 2201-4349 (online) http://dx.doi.org/10.3853/j.2201-4349.68.2016.1669 The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894 (Crustacea, Syncarida, Anaspididae) SHANE T. AHYONG Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia, and School of Biological, Earth & Environmental Sciences, University of New South Wales NSW 2052, Australia [email protected] ABstRACT. The Tasmanian mountain shrimps of the genus Anaspides Thomson, 1894 (Syncarida: Anaspididae), are endemic to Tasmania and often regarded as “living fossils” owing to the retention of numerous putatively plesiomorphic eumalacostracan traits and minimal morphological change since the Triassic. All live in cold, usually high altitude fresh-waters. Until recently, only two species were recognised: the presumed widespread A. tasmaniae (Thomson, 1893) (type species) and A. spinulae Williams, 1965a, from Lake St Clair, whose validity was frequently questioned. Independent morphological and molecular studies revealed previously unrecognized taxonomic diversity, resulting in preliminary descriptions of three new species in 2015. Anaspides is revised based on extensive collections from throughout Tasmania. Telson structure and male secondary sexual characters proved taxonomically instrumental. Seven species are recognized of which two are new to science; all are fully figured and morphological variation is discussed in detail. Rather than being widespread, Anaspides tasmaniae is restricted to Mount Wellington; A. spinulae is a valid species known only from Lake St Clair. Two species, A. clarkei Ahyong, 2015, and A. eberhardi sp. nov. occur only in caves of the Ida Bay-Hastings karst systems and Junee-Florentine systems, respectively. The three widest ranging species (A. jarmani Ahyong, 2015, A. swaini Ahyong, 2015, and A. richardsoni sp. nov.) are primarily epigean and each contains several morphological forms that might warrant further taxonomic subdivision. Distributions of species of Anaspides are largely discrete and broadly correspond to the biogeographical discontinuity known as Tyler’s Line, dividing the drier eastern from the wetter western parts of Tasmania. Caves are believed to have acted as oligothermal refuges for Anaspides in the past, and it is notable that the specimens from the lowest altitudes today are all from caves, including the northernmost record of Anaspides (A. richardsoni, Great Western Cave, Gunns Plain, 109 m asl). Given the revised taxonomy of Anaspides, with significantly altered species distributions, the conservation status of all species of the genus requires review. KEYWORDS. Crustacea; Anaspidacea; Anaspides; Tasmania; freshwater; shrimp. AHYONG, SHANE T. 2016. The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894 (Crustacea, Syncarida, Anaspididae). Records of the Australian Museum 68(7): 313–364. http://dx.doi.org/10.3853/j.2201-4349.68.2016.1669 314 Records of the Australian Museum (2016) Vol. 68 When G. M. Thomson (1893) described Anaspides tasmaniae Washington D.C. (USNM); Universidade Federal de Minas (as Anaspis) from Mt Wellington, Tasmania, he was not Gerais, Brazil (UFMG); Zoological Collection, Universität aware of the significance of his discovery, placing his new Rostock (ZRSO); and Yale Peabody Museum, Connecticut find in the now defunct Schizopoda, alongside mysidacean (YPM). Material examined is listed in approximately north– and euphausiacean shrimps. It was Calman (1897), however, south and east–west order and grouped according to regions who recognized the affinities of the living Anaspides or localities. In the case of “spinose” A. spinulae-like forms with the Palaeozoic fossil syncarids, and would later of A. richardsoni, which occur in the Walls of Jerusalem area recognize a separate eumalacostracan order, Anaspidacea and Western Lakes, they are grouped separately or indicated Calman, 1904 (Brooks, 1962; Schram, 1984). Anaspides is as such within each entry to distinguish them from the typical remarkable because it retains many putatively plesiomorphic non-spinose forms. Similarly, the three morphological forms eumalacostracan traits, such as minimal tagmatisation, of A. richardsoni from the Mole Creek Karst area are also a relatively complete suite of thoracopodal rami, and an grouped for convenience. elementary cardioid escape reaction (Schram & Hessler, 1984). As a result, Anaspides has attracted considerable Taxonomic characters in Anaspides scientific interest as a potential “living fossil” closely resembling its Triassic forbears and possibly reflecting the Species of Anaspides are morphologically conservative, eumalacostracan stem condition (Coineau & Camacho, 2013). exhibiting few differences in major structures. Pleonal Until 2015, only two species of Anaspides were recognised, spination, considered diagnostic of A. spinulae by Williams with A. tasmaniae (Thomson, 1893) (type locality: Mt (1965a), is more variable than previously thought, but when Wellington) accorded a wide range throughout most of considered in combination with other characters, remains central, western and southern Tasmania from both epigean taxonomically useful. Several new taxonomic features are and subterranean habitats, and A. spinulae Williams, 1965a, recognized here and given primacy in separating species— believed to be restricted to central Tasmania from Lake St the shape and spination of the telson, and secondary sexual Clair (type locality) and immediate environs. Additionally, characters of adult males. Anaspides occurs in subterranean habitats throughout The shape and armature of the posterior margin of the Tasmania, with some having troglobitic adaptations such as eye telson, whether evenly rounded or angular, and whether and pigment reduction and unusual telson morphology in which armed with stout, well-spaced spines (Figs 8, 12) or lined the posterior telson spines are few, stout and widely spaced, with numerous finely-set spines (Figs 21, 22), is diagnostic. rather than finely spaced and slender—the “cave type” telson These features are taxonomically effective across a wide (Eberhard et al., 1991; Clarke, 2006). Until recently, taxonomic size range, noting that the posterior margin of the telson in questions surrounding Anaspides focussed on whether A. the earliest post-hatching juveniles has relatively few spines spinulae was distinct from the presumably widespread A. and a median cleft or emargination (Figs 33P, 34T), which tasmaniae, and whether subterranean Anaspides with the rapidly closes with increasing size to form a rounded or “cave-type” telson constituted a separate species (O’Brien, angular spinose margin (Fig. 23B,C). 1990). Preliminary molecular investigations (Jarman & Elliot, Two sexually dimorphic features of males have proven 2000; Andrew, 2005), although not decisive with regards to esp ecially useful: the modified inner antennular flagellum and previous taxonomic questions, nevertheless identified multiple pleopod 1 endopod. In adult males, the inner (= accessory) divergent lineages within Anaspides. Independent, parallel flagellum of the antennule is modified to form a clasping morphological and molecular investigations also recognized structure used to hold females, presumably during courtship significant heterogeneity within the genus, not only among (Hickman, 1937). Rather than being straight and flexible as in epigean, but also subterranean forms, leading to preliminary females and juvenile males, the proximal portion of the adult accounts of three new species of Anaspides (Ahyong, 2015) male inner flagellum (articles 1–8) is semi-rigid and double- in advance of a full taxonomic revision of the genus. The bent into an S-shape, being strongly curled downwardly and present study revises Anaspides, restricting A. tasmaniae to then turning upwards and horizontally in the vicinity of the Mt Wellington, recognizes as valid A. spinulae, and five other sixth and seventh articles (Figs 1, 2D). Each of the proximal species of which two are described as new. eight articles of the modified inner antennular flagellum has several short, curled setae on the dorsal and lateral surfaces, Materials and methods herein called “finger setae”. The proximal 5 articles are as long as or shorter than wide, articles 6 and 7 are more elongated, Measurements of specimens are of total body length, with an obtusely angled inner margin of which the article 7 measured from the apex of the rostrum to the tip of the bears 1–5 slender, prominent, stiff, conical protrusions on the telson. Abbreviations: above sea-level (asl); indeterminate inner margin, herein called clasping spines (Fig. 2D). The (indet); juvenile (juv.); Mount (Mt); station (stn). Almost 3600 number of clasping spines is diagnostic and should always specimens were examined, deposited in the collections of the be examined when making identifications. Both left and right Aust ralian Museum, Sydney (AM); Haswell Museum, Uni- antennules should always be examined in case of damage or versity of Sydney (HMUS); Museum of Zoology, University regeneration from damage to these delicate structures. Note of São Paolo, Brazil (MZUSP); Museum of Comparative also that the clasping spines usually arise sequentially and Zoology, Harvard University (MCZ); Museum Victoria, increase in size with successive moults, so juveniles of A. Melbourne