A new genus and two new species of land snail from geographically remote lithorefugial habitats in Queensland including a remarkable range extension for Quistrachia Iredale, 1939 (Gastropoda: Eupulmonata: Camaenidae) John STANISIC Honorary Research Fellow, Biodiversity Program, Queensland Museum, PO Box 3300, South Brisbane, Qld 4101, Australia. Email: [email protected] Citation: Stanisic, J. 2016. A new genus and two new species of land snail from geographically remote lithorefugial habitats in Queensland, including a remarkable range extension for Quistrachia Iredale, 1939 (Gastropoda: Eupulmonata: Camaenidae). Memoirs of the Queensland Museum-Nature 60: 43-51. Brisbane. ISSN 2204-1478 (Online) ISSN 0079-8835 (Print). Accepted 4 August 2016 First published online: 31 October 2016. https://dx.doi.Org/10.17082/j.2204-1478.60.2016.2016-12 LSID: urn:lsid:zoobank.org:pub:1E91C697-1818-44F5-9F03-28414CFF38A5 ABSTRACT One new genus and two new species of camaenid are described from geographically remote lithorefugial habitats in Queensland. The two habitats have contrasting environments characterised by radically different geology, climate and vegetation. Lorelliana gen. nov. is introduced to accommodate L hoskini sp. nov. from the high rainfall, rainforest peppered, granitic lithorefugia of Cape Melville, Cape York Peninsula. Quistrachia nevbrownlowi sp. nov. is described from the quartzic limestone refugia of the low rainfall, often drought affected, spinifex grassland of Mt Unbunmaroo (= Black Mountain), south-western Queensland. The survival of each species is discussed in relation to the significance of lithorefugia as habitat for land snails in otherwise snail-hostile environments. ¥ new genus, new species, Gastropoda, Camaenidae, lithorefugia. The Camaenidae is a family of land snails lithologies, vegetation and rainfall. In each that is particularly diverse in the rainforests case, adapting to life in lithorefugia has been of eastern Queensland (Stanisic et al. 2010). critical to their survival. Long recognised as a late Miocene invader Lorelliana hoskini gen. et sp. nov. occurs among from land masses to Australia's north the boulder strewn, rainforest habitats of the (Bishop 1981; Solem 1992, 1997), the family Melville Range, Cape York Peninsula. Cape has managed to radiate extensively in many York Peninsula is a bioregion characterised by areas of Australia. In eastern Australia extensive savanna woodlands. Humid tropical camaenids are prominent snails in both the rainforest occurs mainly in the eastern ranges wetter humid coastal rainforests and the drier (Mcllwraith Range, Iron Range), along some subcoastal and inland rainforests. Relatively water courses (e.g. Wenlock River, Jardine few camaenids have made the transition to River) and at the tip of Cape York (Lockerbie much drier eucalypt forests and woodland in Scrub). Elsewhere along the east coast tropical eastern Australia. Greatest diversity occurs in rainforest occurs in remnant scattered pockets the subtropical and tropical rainforests that in gully or rocky headland environments range from central New South Wales to the such as those at Cape Melville. This area is Wet Tropics of northern Queensland. The two subject to high annual rainfall which combined Queensland camaenids described herein occur with the moisture retaining qualities of the in rocky habitats in geographically remote granitic lithorefugia enables it to support environments characterised by differing tropical rainforest communities. Camaenid Memoirs of the Queensland Museum I Nature • 2016 • 60 • www.qm.qld.gov.au 43 Stanisic, J. diversity on Cape York reflects the pattern of SYSTEMATICS scattered, relatively small rainforest patches and consequently is much lower than it is in the vast Infraorder EUPULMONATA rainforest massif of the Wet Tropics.. .but local Superfamily HELICOIDEA endemism is high. L. hoskini sp. nov. is one such endemic species. Family CAMAENIDAE At the other end of the climatic spectrum Genus Lorelliana gen. nov. Quistrachia nevbrownlowi sp. nov. is described from the dry spinifex grasslands of Mt Etymology. Named for Lorelle Holcroft, former Unbunmaroo (= Black Mountain), via Boulia, deputy principal and teacher at Queensland's Samford State School in recognition of her efforts in south-western Queensland (SWQ). Here the science education. annual rainfall is consistently very low in Diagnosis. Shell medium sized, creamy yellow a pattern often compounded by extensive with reddish brown subsutural and peripheral year-long droughts. Q. nevbrownlowi sp. nov. spiral bands, acutely depressedly trochoidal is shown to belong to a genus of land snail with a sharply carinate periphery; protoconch otherwise only present in the Pilbara and smooth, teleoconch mainly smooth with vague southern Kimberley, north-western Western radial growth ridges, base shiny; aperture Australia. This extraordinary disjunct ovately lunate, lip reflected; umbilicus small distribution is unique to the family and is but open. Penis sheath present, thick-walled. probably an artefact of a much wider generic Penis sub-cylindrical, internally with a dense distribution in the past that has been splintered pattern of rhomboidal pilasters, without verge by the continued aridification of the continent or prominent pilaster. Epiphallus long with since the Miocene. The quartzic lithorefugium slender ascending arm, descending arm of Black Mountain would have been critical in thick, folded within penial sheath, entering penis the survival of the species in the face of this through a simple pore; epiphallic caecum present. gradual but dramatic climatic shift. Type species. Lorelliana hoskini sp. nov.-herein designated. MATERIAL AND METHODS Lorelliana hoskini sp. nov. (Figs 1, 2A-C, 3) Material used in this study is held in the collections of the Queensland Museum Etymology. Named for Conrad Hoskin, in recognition (QMMO). Studies of shell characters were of his work on Cape Melville boulder field vertebrates. carried out on specimens in the museum's dry Preferred common name. Cape Melville Talus-snail. collection (RC) and anatomical studies were Material examined. Holotype. QMMO85061, shell based on ethanol preserved samples (SC). RC/animal SC, Cape Melville, Melville Range, FNQ, Measurements of shell characters (height, (14°17/55"S, 144°28'50"E), rainforest on rocky talus, on granite boulders, 10.ii.2013, coll. C. Hoskin, K. diameter, umbilical width) were made using Aland. Height of shell 10.46 mm, diameter 15.84 callipers with a precision of 0.01 mm. Whorl mm, umbilical width 2.29 mm, H/D = 0.66, D/U = 5.68, number of whorls 4.125. counts were made to the nearest 1/8 whorl. Several representatives of the species from their Paratypes. QMMO80774, shell RC/animal SC, same data as holotype; QMM085060, 6RC, Cape Melville, respective type localities were dissected and Melville Range, FNQ, (14°17'55"S, 144°28'50"E), studied using a WILD M5 stereo microscope rainforest on rocky talus, in litter, 15.xii.2013, coll. C. Hoskin, H. Hines. with drawing apparatus in order to confirm stability of reproductive structures. Diagnosis. As for genus. 44 Memoirs of the Queensland Museum I Nature • 2016 • 60 Land snails from geographically remote lithorefugial habitats in Queensland radial growth ridges, base shiny. Aperture ovately lunate, lip reflected, white; umbilicus small but open, umbilical width 2.03-2.66 mm (mean 2.37 mm), D/U = 6.81-8.54 (mean 7.74). Based on eight measured shells (QMMO80774, QMM085060, QMMO85061). Genitalia. Penis sheath (PS) present, thick-walled. Penis (P) sub-cylindrical, internally with a dense pattern of rhomboidal pilasters (PP), without verge or prominent main pilaster. Epiphallus (E) long with slender ascending arm, descending arm thick, folded within penial sheath, entering penis through a simple pore (EP); epiphallic caecum (EC) present. Vas deferens (VD) short. Penial retractor muscle (PRM) inserted on the epiphallus at the point of reflexion. Vagina (V) short, less than half length of penis, internally with longitudinal thickenings. Atrium (Y) simple. Free oviduct (UV) almost equal in length to vagina; bursa copulatrix (not figured) with a relatively wide stalk (BC) located at the base of the albumen gland; prostate (DG) and uterus FIG. 1. Lorelliana hoskini gen. et sp. nov. QMMO85061, (UT) without unusual features. Albumen holotype. Genitalia. Scale line = 10 mm. gland, hermaphroditic duct and ovotestes not illustrated. Based on two dissected adults Description. Shell medium sized, creamy yellow (QMMO80774, QMMO85061). with reddish brown subsutural and peripheral Animal. Animal (Fig. 3) orange with orange foot, spiral bands, acutely depressedly trochoidal black head and neck, upper and lower tentacles with a sharply carinate periphery, whorls 4.125- orange. Pallial region with black pallial collar 4.75 (mean 4.5), the last descending slightly in and two black stripes running rearwards and front; spire slightly elevated. Height of shell located on either side of the pallial roof. Head 7.67-8.44 mm (mean 8.11 mm), diameter of shell wart present. 17.34-18.80 mm (mean 18.10 mm), H/D = 0.43- 0.47 (mean 0.45). Protoconch of 1% whorls, Habitat and ecology. Scattered patches of rainforest shiny, teleoconch mainly smooth with vague on granite talus; living among granite boulders. FIG. 2. Lorelliana hoskini gen. et sp. nov. QMMO85061, holotype. Shell views. A, apertural; B, dorsal; C, ventral. Scale lines = 10 mm. Memoirs of the Queensland Museum I Nature • 2016 • 60 45 Stanisic, J. different shell sculpture and the contrasting anatomical details (Solem 1989). Genus Qnistrachia Iredale, 1939 Diagnosis. See Solem (1997). Type species. Trachia monogramma Ancey, 1898-by original designation. Qnistrachia nevbrownlowi sp. nov. (Figs 5A, B; 6A-C) Etymology. Named for Neville Brownlow, Director of Earthborn Australia Pty Ltd, in recognition of his efforts in organic waste recycling. Preferred common name. Boulia Globular Snail. Material examined. Holotype. QMMO85063, shell RC/animal SC, Boulia, c. 50 km NE at Black Mt, SWQ, (22°33/03/,S, 140°13'22"E), along creek in flood debris, 20.xi.2002, coll. R. Crookshanks. Height of shell 10.46 mm, diameter 15.84 mm, umbilical width 2.29 mm, H/D = 0.66, D/U = 5.68, number of whorls 4.625. FIG. 3. Lorelliana hoskini gen. et sp. nov. Live animal. Paratypes. QMMO71806,3SC, same data as holotype; QMMO71805, 4 SC, Boulia, c. 50 km NE at Black Mt, SWQ, (22°36,10"S, 140°13,32"E), along creek Distribution. Known only from the type locality. in flood debris, 20.xi.2002, coll. R. Crookshanks; QMM072256, 6SC, Boulia, c. 50 km NE at Black Mt, Remarks. Lorelliana hoskini sp. nov. is endemic SWQ, (22°38'27"S, 140°16T4/,E), along creek in flood to the Melville Range boulder fields (Fig. 4) and debris, 20.xi.2002, coll. R. Crookshanks; QMMO71807, 3SC/1RC, Boulia, c. 50 km NE at Black Mt, SWQ, can be readily distinguished by the combination (22°30'00"S, 140°16'30"E), along dry gully under dead of sharply carinate, very depressed, trochoidal spinifex, 20.xi.2002, coll. R. Crookshanks; QMM04995, shell and the orange animal with orange 50+ RC, AMSC.517369,6RC; WAMS70841,6RC; Black tentacles and black head. The relationships of Mt, Warenda Station, c. 60 km NE Boulia, gibber and spinifex plain/limestone, on ground, alt. c. 180 m, the species are yet to be determined however, viii.1971, coll. A. Hiller, M. Wade. the short penis with a thick sheath, internal wall sculpture of dense rhomboidal pilasters, no Other material. QMMO15053, 7SC, Black Mt, NE Boulia, WCQ, limestone area, viii.1984, coll. A. verge and no main pilaster combined with an Rozefelds; QMMO85064, 4RC, locality given only epiphallus which is partially folded within the as Queensland (= Black Mountain area, via Boulia), penial sheath are a combination of anatomical SWQ, 1975, coll. M. A. Archer. characters which separate this species from all Diagnosis. Shell medium-sized, with reddish other Cape York Peninsula camaenids thus far brown suffusion on upper whorl surface dissected (Stanisic, unpubl.). extending below whorl periphery and a darker, Interestingly the shell of L. hoskini bears an narrow subsutural band and less prominent uncanny resemblance to that of Trochomorpha peripheral band, base white; helicoid with an melvillensis Solem, 1989 (Family Trochomorphidae) elevated spire and rounded whorls; teleoconch which curiously comes from a somewhat sculpture of closely spaced, anastomosing similarly named locality in Melville Island radial ridges; aperture roundly lunate, lip white, (Tiwi Islands), Northern Territory. FFowever, the expanded and reflected; umbilicus narrowly superficial conchological resemblance between open. Penis with a thick sheath, internally with the two species is not reflected in vastly a main long, bifurcated pilaster, penis wall with 46 Memoirs of the Queensland Museum I Nature • 2016 • 60 Land snails from geographically remote lithorefugial habitats in Queensland FIG. 4. Cape Melville boulders and rainforest. Image: Conrad Hoskin. a series of longitudinally arranged diamond¬ H/D = 0.58-0.69 (mean 0.65). Protoconch of shaped pustules merging into longitudinal IV2 whorls sculptured with irregular radial pilasters basally. Vas deferens entering penis ridges, teleoconch sculpture of closely spaced, sheath basally and ascending almost entire anastomosing radial ridges extending onto length of penis, epiphallus short, entering penis base. Aperture roundly lunate, lip white, apically through a simple pore; bursa copulatrix expanded and reflected; umbilicus narrowly with a short stalk and expanded head located at open, umbilical width 2.10-2.78 mm (mean 2.46 the base of the prostate-uterus. mm), D/U = 4.82-7.17 (mean 6.03). Based on 22 measured adults (QMM04995, QMMO71807, Description. Shell medium-sized, with reddish QMMO85063). brown suffusion on upper whorl surface extending below whorl periphery and a darker, Genitalia. Penis (P) with a thick sheath (PS), narrow reddish brown subsutural band and internally with main pilaster bifurcated (PPM), less prominent reddish brown peripheral band, penis wall with a series of longitudinally base white; helicoid with an elevated spire and arranged diamond-shaped pustules (PP) 4.125-4.625 (mean 4.5-) rounded whorls, the last merging into longitudinal pilasters basally. Vas descending slowly in front; spire moderately deferens (VD) entering penis sheath basally elevated. Height of shell 7.96-11.43 mm (mean 9.62 (VDE) and ascending almost entire length of mm), diameter 13.15-16.94 mm (mean 14.72 mm). penis. Epiphallus (E) short, entering penis Memoirs of the Queensland Museum I Nature • 2016 • 60 47 Stanisic, J. FIG. 5. Quistrachia nevbrownlowi sp. nov. QMMO71807, paratype. A, genitalia. Scale lines = 5 mm; B, penial complex. Scale lines = 2.5 mm. apically through a simple pore (EP). Vagina Habitat and ecology. Spinifex on quartzic (V) equal in length to the penis, internally with limestone; living under spinifex, rocks and longitudinal thickenings, free oviduct (UV) scattered debris. relatively long; bursa copulatrix (BC) short with Distribution. Known only from Mt Unbunmaroo an expanded head located at the base of the (= Black Mountain), NE Boulia, SWQ. prostate-uterus; prostate (DG) and uterus (UT) without unusual features. Albumen gland (GG) Remarks. Quistrachia nevbrownlowi sp. nov. slender, hermaphroditic (GD) duct typical. has only been recorded in the vicinity of Mt Animal. Preserved animal with light-brown body, Unbunmaroo (= Black Mountain), via Boulia, head region slightly darker. Head wart absent. SWQ. The habitat is one of quartzic limestone FIG. 6. Quistrachia nevbrownlowi sp. nov. QMMO85063, holotype. Shell views. A, apertural; B, dorsal; C, ventral. Scale lines = 10 mm. 148 Memoirs of the Queensland Museum I Nature • 2016 • 60 Land snails from geographically remote lithorefugial habitats in Queensland TABLE 1. Local variation in shells of Lorelliana hoskini sp. nov. and Quistrachia nevbrownlowi sp. nov. Taxon N Diameter Height Umbilical H/D ratio D/U ratio Number (mm) (mm) width (mm) of whorls L. hoskini sp. nov. QMMO85061 holotype 1 17.34 7.67 2.03 0.44 8.54 4.125 QMMO80774 paratype 1 18.80 8.44 2.36 0.45 7.97 4.375 QMM085060 paratypes 6 17.52-18.80 7.69-8.36 2.23-2.66 0.43-0.47 6.81-8.43 4.5-4.75 Q. nevbrownlowi sp. nov. QMMO85063 holotype 1 16.59 10.65 2.79 0.64 5.95 4.625 QMM04995 paratypes 20 13.15-16.94 7.96-11.45 2.10-2.78 0.58-0.69 4.82-7.17 4.125-4.625 QMMO71807 paratype 1 15.84 10.46 2.59 0.66 6.12 4.5 and spinifex grassland (Fig. 7). Q. nevbrownlowi nov. these land snails have displayed resilience is the sole outlier of a genus whose main in the face of climatically induced, snail-hostile mass of species (9) is located in the Pilbara, environmental change. Dampierland and southern Kimberley of Lorelliana hoskini sp. nov. lives among the north-western Western Australia. The basal granite talus of Cape Melville in an area where entrance of the vas deferens into the penial rainforest has contracted and now occurs in sheath, poorly defined epiphallus, bifurcated gullies and on elevated plateaux within the main penial pilaster and short bursa copulatrix boulder fields of the greater Melville Range, are all features which clearly place the species Cape York Peninsula (Fig. 8). Rainforest in in the otherwise Western Australian genus. The eastern Australia receded into mountain top genital features most closely ally Q. nevbrownlowi refugia and along moist drainage lines from to the Pilbara species Q. legendrei Solem, 1997. the mid-Miocene onwards, and with greater Quistrachia nevbrownlowi was noted by Solem amplitude and rapidity during the Quaternary (1997:1876-1877) in his revision of Quistrachia (Galloway & Kemp 1981). Deeply layered but never described. rocky outcrops such as those in the Melville Range would have provided persistent moist DISCUSSION and cool microhabitats for mesic organisms such as land snails (Couper & Hoskin 2008). The present study highlights the importance Having a shell for protection and one which of lithorefugia for the survival of land snails in is designed for life in crevices (flattened) two contrasting Queensland environments. The combined with a nocturnal habit would have granite boulder fields of the Melville Range, pre-adapted the species for survival in such Cape York Peninsula is in an area of relatively a specialised environment. Persistently high high annual rainfall (average annual rainfall rainfall and shelter from fire would also have > 1500 mm). The quartzic limestones of Mt been major contributing factors. The area is Unbunmaroo (= Black Mountain) on the other already remarkable for the number of endemic hand have very low rainfall (mean < 250 mm vertebrate species (three frogs, two skinks and per annum) with sporadic episodes of multi¬ one gecko) all of which have connections with year droughts. In the case of both Lorelliana the rainforests of the Wet Tropics further south hoskini sp. nov. and Quistrachia nevbrownlowi sp (Hoskin & Couper 2013; Hoskin 2013a, b, 2014). Memoirs of the Queensland Museum I Nature • 2016 • 60 49 Stanisic, J. FIG. 7. The spinifex covered terrain around Mount Unbunmaroo (= Black Mountain), via Boulia, SWQ. Image: Graham Harrington. L. hoskini is the first endemic invertebrate thus far recorded from the locality. Its relationships are unknown at this time but may also lie within the Wet Tropics where much of the land snail fauna has yet to be collected let alone described. Quistrachia nevbrownlowi sp. nov. lives among the limestone and spinifex of the Mt Unbunmaroo (= Black Mountain) area, via Boulia, SWQ. However, its nearest relatives are almost 2000 km away on the western edge of the continent, usually in the rocky talus strewn spinifex grasslands of the Pilbara and Kimberley of north-western Western Australia (Fig. 8). While some dry adapted pupillids bridge this geographic gap (Glyptopupoides egregia, Gastrocopta spp.) the distribution pattern is quite unusual in the context of the Camaenidae. There are camaenid genera such as Torresitrachia and Xanthomelon with so-called trans-Australian distributions FIG. 8. Distribution map. Lorelliana hoskini gen. et (Solem 1979), but the range extension of a sp. nov. (star): Cape Melville, north Queensland; camaenid genus from its core in the Pilbara- Quistrachia nevbrownlowi sp. nov. (triangle): Kimberley to a disjunct species in south¬ Mt Unbunmaroo, SWQ; Other Quistrachia spp., north-western Western Australia: 1, Pilbara; western Queensland, without any intervening 2, Dampierland and southern Kimberley. congeners, is somewhat remarkable. 50 Memoirs of the Queensland Museum I Nature • 2016 • 60 Land snails from geographically remote lithorefugial habitats in Queensland The extraordinary range disjunction between LITERATURE CITED Q. nevbrownlowi and the main mass of Quistrachia Bishop, M.J. 1981. The biogeography and evolution species implies that there was once some of Australian land snails. Pp. 924-954. In, Keast, historical connection of Quistrachia habitat A. (ed.). Ecological biogeography of Australia. (Dr W. between the two regions. It is reasonable to Junk: the Hague) (http://dx.doi.org/10.1007/978- speculate that intervening species would 94-009-8629-9_32). have become extinct along with this historical Couper, P.J. & Hoskin, C.J. 2008. Litho-refugia: the habitat through climatic attrition in the face importance of rock landscapes for the long¬ term persistence of Australian rainforest fauna. of widespread continental aridification. The Australian Zoologist 34: 554-560 (http://dx.doi. onset of aridity was accompanied by extensive org/10.1007/ 978-94-009-8629-9_32). desertification in central areas of the continent Galloway, R.W. & Kemp, E.M. 1981. Late Cainozoic whereby sand dunes covered much of the environments in Australia. Pp. 52-80. In, Keast, region. Survival of present day Quistrachia A. (ed.). Ecological biogeography of Australia. species relied on their ability to show resilience (Dr W. Junk: The Hague) (http://dx.doi. in the face of this aridification and adapt to life org/10.1007/ 978-94-009-8629-9_4). in scattered rock piles throughout the historical Hoskin, C.J. 2013a. A new frog species (Microhylidae: range. The availability of moisture and shelter Cophixalus) from boulder-pile habitat of Cape Melville, north-east Australia. Zootaxa 3722: 61-72 sites among the rocky talus and the role of rock (http: / / dx.doi.org/10.11646/ zootaxa.3722.1.5). outcrops in excluding fire are all key attributes Hoskin, C.J. 2013b. A new skink (Scincidae: beneficial for the continued survival of land snails. Saproscincus) from rocky rainforest habitat on That the current Western Australian species Cape Melville, north-east Australia. Zootaxa now live under rocks and litter among rock 3722: 385-395 (http://dx.doi.org/10.11646/ outcrops in the climatically challenging Pilbara zootaxa.3722.3.7). and Kimberley is practical evidence for such a Hoskin, C.J. & Couper, P. 2013. A spectacular new leaf¬ proposition. In an area of western Queensland tailed gecko (Carphodactylidae: Saltuarius) from that also endures unforgiving climatic conditions the Melville Range, north-east Australia. Zootaxa 3717: 543-588 (http://dx.doi.org/10.11646/ the presence of the limestone lithorefugium that zootaxa.3717.4.6). is Black Mountain probably played a major Hoskin, C.J. 2014. A new skink (Scincidae: Carlia) from role in the survival of Q. nevbrownlowi. the rainforest uplands of Cape Melville, north-east In conclusion, it should be added that this study Australia. Zootaxa 3869: 224-236 (http://dx.doi. org/10.11646/ zootaxa.3869.3.2). once again underscores the part that land snails (and other invertebrates) can play in identifying Solem, A. 1979. Camaenid land snails from western and central Australia (Mollusca: Pulmonata: places of environmental significance. These Camaenidae). I. Taxa with trans-Australian organisms are too often ignored in environmental distribution. Records of the Western Australian survey and assessment. Yet they have the innate Museum, Supplement 10: 5-142. ability to identify special habitats in their roles as Solem, A. 1992. Camaenid land snails from southern both environmental indicators and biodiversity and eastern Australia, excluding Kangaroo predictors, in particular the moisture loving Island. Records of the South Australian Museum land snails. Monograph Series 2:1-425. Solem, A. 1997. Camaenid land snails from Western and central Australia (Mollusca: Pulmonata: ACKNOWLEDGEMENTS Camaenidae). VII. Taxa from Dampierland through the Nullarbor. Records of the Western I wish to particularly thank Conrad Hoskin Australian Museum, Supplement 50:1461-1906. (James Cook University, Townsville, Queensland) Stanisic, J., Shea, M., Potter, D. & Griffiths, O. 2010. and Kieran Aland (Queensland Museum, Australian land snails Volume 1: A Field guide Brisbane) for their efforts in making material of the to eastern Australian species. (Bioculture Press: Cape Melville Talus-snail available for this study. Mauritius). 596pp. Memoirs of the Queensland Museum I Nature • 2016 • 60 51