Journal of the Royal Society of Western Australia
ISSN 0035-922X CONTENTS
Page Hydrological change escalates risk of ecosystem stress in Australia's threatened biodiversity hotspot P Horwitz, D Bradshaw, S Hopper, P Davies, R Froend & F Bradshaw
Origins and nature of vessels in monocotyledons. 10. Boryaceae: xeromorphic xylem structure in a resurrection plant S Carlquist, E L Schneider & K Kenneally 13 Greater Bilby (Macrotis lagotis) burrows, diggings and scats in the Pilbara G G Thompson & S A Thompson 21 Age estimates of Sporolithon durum (Corallinales, Rhodophyta) from Rottnest Island, Western Australia, based on radiocarbon-dating methods N A Goldberg & J N Heine 27 A new species of soldier crab, Mictyris occidentalis (Crustacea: Decapoda: Brachyura: Mictyridae) from Western Australia,with congener comparisons J Unno 31 Range extension for the Black-headed Worm-lizard, Aprasia picturata (Squamata: Pygopodidae) S A Thompson & G G Thompson 51
Volume 91 Part 1 March 2008 The Royal Society of Western Australia
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Hydrological change escalates risk of ecosystem stress in threatened biodiversity hotspot
P Horwitz1*, D Bradshaw2, S Hopper3, P Davies4, R Froend1 and F Bradshaw2 it 1 School of Natural Sciences, Edith Cowan University, Perth, Australia 2School of Animal Biology and Centre for Native Animal Research, University of Western Australia, Perth, 6009, Australia 3Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK and School of Plant Biology, University of Western Australia, Perth, 6009, Australia 4 Centre of Excellence in Natural Resource Management, University of Western Australia, Albany, 6330, Australia
* To whom correspondence should be addressed. E3 [email protected]
Manuscript received August 2007; accepted February 2008
Abstract
The southwestern comer of the Australian continent has been identified as a global "biodiversity hotspot", defined as an area where "exceptional concentrations of endemic species are undergoing exceptional loss of habitat". In this paper we reconsider the reasons for this conservation priority. We briefly review significant characteristics of the flora and fauna, and the way threatening processes are escalating ecosystem stress to these conservation values. Our specific aim is to examine the ecological consequences of hydrological change, including emergent issues such as climate change, and focus on the coastal plains in higher rainfall zones where the majority of the Western Australian population resides. Here we argue that human-driven and/or climatically- driven hydrological change deserve greater attention, since they: i) directly escalate the risk of extinction for some components of the biota, or ii) are underlying and/or contributing factors in the manifestation of other threats to the biota, and may complicate or exacerbate some of those threats (such as fire, Phytophthora and the spread of weeds). This paper briefly outlines the challenges to the region's biodiversity posed by hydrological change. We suggest a societal adoption of approaches based on water literacy will be necessary to avoid irreversible changes associated with a continued reliance on water resource developments and other energy/water intensive industrial activities.
Keywords: biodiversity, hotspot, conservation, SWAFR, land clearing, salinity, fire, Phytophthora, climate change, groundwater extraction, water literacy
Introduction Malcolm et al. (2006) assessed the 25 biodiversity hotspots using major vegetation types as proxies for The southwestern corner of the Australian continent natural habitats, and identified southwest Australia as has been identified as a global "biodiversity hotspot", one of six especially-vulnerable regions. defined as an area where "exceptional concentrations of endemic species are undergoing exceptional loss of The biodiversity hotspot described in this paper habitat" (Myers et al. 2000, p. 853). Twenty five such coincides approximately with the South-West Botanical threatened hotspots were identified in 2000, and the Province (Beard 1980; Beard 1999), updated to the number has since been increased to 34 however only one Southwest Australian Floristic Region (SWAFR, sensu occurs in Australia (Mittermeier et al. 2004). The original Hopper & Gioia, 2004). It extends from forested intention of these determinations has been to raise ecosystems dominated by large eucalypts and coastal awareness, and encourage conservation planners to peatlands, heathlands and fresh perennial wetland prioritise their investment in mitigating threatening systems in the extreme southwest, to species-rich processes. We note that in Western Australia the kwongan sandplain heaths in the drier northern and identification of the southwestern part of the State as a eastern regions between Shark Bay and Israelite Bay, and global priority has too-often been one of celebrating inland to eucalypt-dominated woodlands and shrublands international recognition for biodiversity rather than a often called the 'wheatbelt' and mallee regions (see prompt to reorientate acitivities to reverse the exceptional Figure 1). It is often described as an 'island' isolated by loss of habitat. The identification of biodiversity hotspots ocean to the west and south, and arid areas to the north has also allowed researchers to undertake coarse-scale and east, spanning average annual rainfalls of over 1200 analyses of emerging threats such as global warming: mm close to the southwestern coast to around 300 mm per annum in the interior. Superimposed on these characteristics is a rapidly-expanding human population concentrated along the coastal margins, where rainfall © Royal Society of Western Australia 2008 has tended to be higher and less variable.
1 Journal of the Royal Society of Western Australia, 91(1), March 2008
Figure 1. Showing the South West Australian Floristic Region (SWAFR), the location of the Gnangara Mound area (on the Swan Coastal Plain), the middle and lower reaches of the Blackwood River, place names mentioned in the text, and Australian bioregions (where GS= Geraldton Sandplains, YAL=Yalgoo, AW= Avon Wheatbelt, WAR= Warren, SWC = Swan Coastal Plain, ESP= Esperance, and JF= Jarrah Forest). Note: the bioregion of Yalgoo was not recognised as part of the SWAFR by Hopper and Gioia (2004) but is included here because it is regarded as part of the southwestern Australian biodiversity hotspot.
In line with Myers et al.'s (2000) intention, in this being endemic (Beard et al. 2000). While representing paper we reconsider the reasons for the conservation only about 0.2% of the Earth's surface in area, this priority of southwestern Australia. We commence by diversity of plant life represents 1% of the world's total briefly reviewing significant characteristics of the flora number of plant species (Hopper & Gioia 2004), and and fauna, and the way threatening processes may points to the global significance of the region from a impact on these conservation values. Our aim is to biodiversity perspective. This rich biodiversity of the examine the ecological consequences of hydrological SWAFR is primarily expressed amongst its angiosperms, change, including emerging issues such as climate especially woody families such as Myrtaceae (1283 change, and particularly focus on the coastal plains in species/subspecies), Proteaceae (859), Fabaceae (540) and higher rainfall zones where the majority of the State's Mimosaceae (503). Other species-rich families include population resides. Here we argue that hydrological Orchidaceae (374) Ericaceae (including Epacridaceae, change directly escalates the risk of extinction for some 297), Asteraceae (280), Goodeniaceae (207), Cyperaceae components of the biota, or is an underlying and/or (199) and Stylidaceae (178) (Hopper & Gioia 2004). contributing cause for other threats to the biota, and may By March 2006, 351 plant species within the region be exacerbated by some of those threats. had been listed as threatened, and approximately 2,500 were of conservation concern according to the State's Biological Characteristics of the Southwest Australian Department of Environment and Conservation Floristic Region threatened species database. Threatened taxa are mostly Estimates of the number of native plant species woody perennials, one third of which are short-lived identified in the region vary from 5,469 (Myers et al. disturbance opportunists and species that typically seed 2000) to 6,759 (Hopper & Gioia 2004) with 49.0% to 53.9% after fire such as Acacia and Grevillea spp. (Atkins 1998;
2 Horwitz et al: Hydrological change stress in a biodiversity hotspot
Bell 2001; Hopper et al. 1990). Perennial herbs figure has been the recognition of a new order of tiny annuals prominently amongst the remaining threatened taxa, of temporary seasonal wetlands, the Hydatellales, more than half being orchids. Spring flowering occurs in comprising a single family Hydatellaceae of two genera, two-thirds of the threatened taxa and 40% possess Hydatella and Trithuria (Saarela et al. 2007). Hydatellales flowers that are probably pollinated by birds and are richest in the SWAFR, but extend to temperate mammals, rather than by insects or wind (Brown et al. eastern Australia, New Zealand and India. Previously 1997; Hopper & Burbidge 1986; Keighery 1982). believed to be obscure monocots, DNA evidence now Invertebrate species also show high levels of regional unequivocally shows that they are among the earliest endemism, with an emphasis on Gondwanan affinities flowering plant lineages, sister to the water lilies (Edward 1989; Main 1996, 1999). While many (Nymphaeales), originating even earlier than invertebrate groups have been inadequately researched, Dasypogonales. there is sufficient evidence to support generalisations regarding patterns of high endemicity: some invertebrate Recognised Threatening Processes groups are associated with particular habitats (such as In their review of biodiversity hotspots, Mittermeier et cave systems (Jasinska et al. 1996) and springs (Knott & al. (2004) regarded habitat destruction, due to clearing of Jasinska 1998), and/or pockets of year-round moisture; native vegetation for agriculture, as being the most the terrestrial isopods are a good example (Judd 2005)); significant historical process to have led to a loss in others have particular relationships with plants or plant biodiversity in the SWAFR. Their principal source of communities (Majer et al. 2001) or have speciated locally information was vegetation mapping conducted for (for example the non-biting midges Chironomidae Western Australia by Shepherd et al. (2002). These latter (Edward 1989)), while others have a local affinity for authors claim, inter alia, that "A total of 119 associations freshwater (Horwitz 1997; Pinder et al. 2004). have been reduced to below 30 per cent of their pre- The vertebrate fauna of the southwest is rich, with 456 European extent and of these, 48 have less than or equal species listed (Myers, et al. 2000), 100 of which are to 10 per cent remaining and two are presumed extinct'. endemic to the ecoregion (How ef al. 1987). These include This clearing of native vegetation in the SWAFR eight species of freshwater fish (Morgan et al. 1998), 24 commenced with the arrival of European settlers in 1829 amphibians, 50 reptiles, 19 birds, and seven mammals. and grew steadily in the early part of the 20lh century Five bird species are listed as threatened under the (Bolton 1972). It accelerated dramatically in the 1960s Commonwealth's Environmental Protection and with the Government's policy of clearing 'a million acres Biodiversity Conservation Act 1999: Carnaby's Black a year' for wheat and sheep farming in the kwongan cockatoo, Baudin's Black cockatoo, the Noisy Scrub-bird sandplain areas bordering Mount Lesueur, and in the (Atrichornis clamosus), the Western whipbird and the region between Albany and Esperance. Beard (1999) Western bristlebird and a number of species of frogs, estimated that the original habitat area of approximately reptiles and fish are also listed. Burbidge and McKenzie 357 km2 in southwest WA has now been reduced by 70% (1989) noted that the extent of extinction of mammals in to 107 km2. Such habitat destruction and alienation has Australia since European settlement has no recent been identified by fauna specialists as the single most parallel on any other continent and southwestern important factor challenging the long-term survival of Australia has experienced one of the greatest levels of threatened and endangered species (McDonald et al. regional extinction of mammals anywhere in Australia 2003). (Short 2004; Woinarski & Braithwaite 1990). Associated with the clearing of native vegetation for However, although the richness of the southwest agriculture have been increasing salinity in waterways region has been recognised at a global level, much of this (Schofield 1990; Bunn & Davies 1992), and the spread of area has still to be systematically surveyed and feral predators, competitors and weeds. Salt brought to catalogued. The Noisy-Scrub bird, Atrichornis clamosus, the surface from rising groundwater contributes to saline was "rediscovered" at Two People Bay, east of Albany in runoff into river systems that were previously fresh. 1961 (Smith 1985) and the discovery of new genera in Surface flow regulation is also extensive and typically recent years, such as the Sunset frog, Spicospina associated with a decline in freshwater biodiversity (Kite flammocaerulea, in 1994 (Roberts et al. 1997), and the et al. 1997). The impact of exotic predators, such as foxes long-thought-to-be extinct Gilbert's potoroo, Potorous and cats, and exotic competitors such as rabbits, sheep gilbertii, also at Two-People Bay (Sinclair et al. 1997), and cattle on species extinction is difficult to quantify but should caution against concluding that the biodiversity universally acknowledged as a major factor in the decline of this important region is well-known (Main & Main of, especially mammalian, populations (Marshall 1966; 1991), Short 2004; Short et al. 1992). Mittermeier et al. (2004) This is the case even more so for non-vascular plants, also regarded the spread of the plant pathogen ("dieback") Phytophthora cinnamomi, as the most fungi (see May 2002) and other microbes, and even for serious current threat: elsewhere it has been dubbed the vascular plants; a third of the 6,759 species known from 'biological bulldozer' (Shearer 1994; Shearer et al. 2004). the SWAFR were described for the first time in the past Phytophthora cinnamomi is listed under the EPBC Act three decades, and at least another 1000 are estimated to remain unnamed (Hopper & Gioia 2004). Moreover, 1999 as "a key threatening process to Australia's biodiversity" and of the plant species in the SWAFR, 40% DNA sequence studies have revealed unexpected are susceptible and 14% highly susceptible to the lineages of great antiquity. Dasypogonales (the world's pathogen (Shearer et al. 2004). most localised order of Angiosperms) has an estimated age of origin of about 120 million years BP, early in the Australia-wide, Cork et al. (2006) used expert opinion Cretaceous. Perhaps the most surprising recent discovery to rank eight pressures impacting on different aspects of
3 Journal of the Royal Society of Western Australia, 91(1), March 2008
biodiversity: total grazing, feral animals, weeds, changed Swan Coastal Plain is a good example where underlying fire regime, habitat fragmentation, vegetation clearing, or distal causes of hydrological changes must include all changed hydrology and salinity. They argue that some of of the following, to various degrees: these drivers will decline due to adequate control (such I. Climate variability: in particular inter-annual as that imposed on land clearing, thereby reducing variability in the pattern of rainfall, temperature habitat-modification pressures); some will continue in the regimes (air and water), cloud cover, wind future, and some (such as climate change) will increase direction and strength, and pan-evaporation in importance and severity. They also argued, inter alia, rates. that "Increasing direct effects on ivetlands and aquatic dependent biodiversity are likely to be made worse with II. Patterns of land-use change: shifts from climate change. Inadequate attention has been paid to woodlands or heath to agriculture and groundwater-dependent biodiversity." horticulture, in some places followed by urban development. For urban areas, land use patterns Burgman et al. (2007) explored threat syndromes and change with age as suburbs become more conservation strategies for Australian plants, finding that vegetated. Each of these shifts in land use is land clearance for agriculture (grazing and cropping) and accompanied by different patterns of surface urbanization have been the primary causes of range runoff and groundwater recharge from rainfall, contractions and habitat loss in the past, responsible for and different types of microclimatic feedbacks to the current status of the majority of threatened Australian the macroclimate. plants. They found that threats growing in importance include disease, salinity, invasive species and changed III. Patterns of water regulation: damming of disturbance regimes. Many species are subject to catchments has substantially reduced flow from common, landscape-level threats that often interact as the Darling Scarp onto the Swan Coastal Plain; threat syndromes. In some cases, effective mitigation nearly 120 years of draining has virtually requires 'simple, low-cost changes in policy, such as eliminated previously extensive areas of more stringent controls on land clearance, strategic fire inundation of the Swan Coastal Plain (see for management, and firmer control on the importation of example Bradby 1997). plant species. Other factors will require greater effort and IV. Patterns of groundwater extraction: intensive new strategies to mitigate, including social and legal localised extractions for horticulture, agriculture initiatives in urban landscapes and broad strategies for and domestic water supplies plus the extraction pathogens, climate change and other landscape-level of water from private residential bores (see processes.' below). The Western Australian Government's most recent V. Water infrastructure: patterns of groundwater State of the Environment Report (Government of Western recharge and behaviour in terms of Australia, 2007) listed 34 environmental issues of priority overconsumptive water use are influenced by for the State. Of these, the top priorities were regarded as inward bound reticulated distribution of potable climate change, population and consumption, water through the Integrated Water Supply greenhouse gas emissions, land salinisation, salinisation Scheme (IWSS), and outward bound collection in of inland waters, introduced animals, weeds, and 'wastewater' facilities before most of it is Phytophthora dieback; all were considered likely to have discharged off-shore. Stormwater runoff is also resulted in on-going deterioration over the last ten years. 'collected' resulting in similar discharge or For inland waters, altered water regimes, loss or concentrated recharge in sumps or wetlands degradation of wetlands, and loss or degradation of (creating a local mounding affect in places; fringing and instream vegetation were listed as Appleyard et al. 1999) belonging to the next most urgent priorities. Within the VI. External drivers: the macroeconomy, lack of SWAFR, the Swan Coastal Plain was shown to be the adequate water pricing, political ideologies and most profoundly affected bioregion (where more than 26 population growth Scenarios influence all of the of the State's 34 environmental issues are "active"). Here, above. intensification of land clearing and continuous development of the coastal margins of the southwest for Hennessy et al. (2007) refer explicitly to southwestern housing, recreation and tourism is coupled with the Australia for actual and future climate change, State's vigorous resources boom, placing on-going particularly increased temperatures and less rainfall. pressure on conservation values with significant Indian Ocean Climate Initiative (2002) demonstrated this alteration to water and nutrient cycles. actuality: average May-October and May-July rainfall in the southwest over the last 25 years has been only 85- These latter arguments suggest that a heightened 90% of the preceding 50-year average and stream inflow attentiveness to hydrological change, the declining water into Perth's reservoirs over the period 1997-2004 is now availability to ecosystems in the populated coastal only 54% of levels recorded from 1975-1996 (see Figure margins and Swan Coastal Plain bioregion in particular, 2). Nicholls (2006) concluded that the rainfall decrease in is warranted. southwest Western Australia is likely due to a combination of increased greenhouse gas concentrations, Climate change and hydrological change natural climate variability and land-use change (building Our desire to separate causes of environmental on observations of atmospheric differentials described by degradation ('threats') is problematic when complex Lyons (2002) between cleared land and uncleared land, systemic factors are at play. Hydrological change on the and modelling work of others).
4 Horwitz et al: Hydrological change stress in a biodiversity hotspot
SWWA rainfall (SW of line between 30S, 11SE ft 35S, 120E)
Year
j -May-October -May-Jufy -August-October
Figure 2. Trends in average yearly rainfall (mm) in the southwest of Western Australia (southwest of a line connecting 30S, 115E & 35S, 120E) from 1952 to 2002 with May-October in blue, May-July in red and August-October in green (from Indian Ocean Climate Initiative 2002).
Perth, the expanding urban area of the Swan Coastal Hydrological change as a threatening process Plain, ranks among those cities most likely to experience Broad ecological linkages between hydrology and future water stress (Flannery 2003; Jenerette & Larsen biology for inland aquatic and terrestrial species, 2006). This is due to the combined influences above, with communities and/or ecosystems in coastal margins of the sustained per-capita water consumption, reduced SWAFR are given in Table 1. The Table shows the range availability of water, and consequent demand responses. of eight possible types of linkages and examples of each. The most accessible groundwater for Perth is that found The scheme is based on the principle that linkage in the superficial sediments, as unconfined aquifers in between hydrology and biology is a question of degree the sands of the coastal plains. The most significant of (similar to Hatton and Evans' (1998) principal criterion these unconfincd aquifers, the Gnangara Mound, occurs that the degree of 'groundwater dependence' is under the northern suburbs of Perth and beyond. The proportional to the fraction of the annual water budget exact contribution that this aquifer makes to Perth's that ecosystem derives from groundwater). The eight water consumption varies from year to year and types are separated primarily along an axis of water according to how the figures are calculated but it is permanence, but secondarily according to the nature of substantial; for instance, at least 60% of IWSS water for the linkage, related to life history characteristics of the Perth metropolitan area comes from the Gnangara organisms and biogeochemical requirements. They Mound, and the more confined, deepdr aquifers that therefore build on the categories of groundwater underlie it (like the Leedcrville Formation and the dependence defined by Hatton and Evans (1998) and Yarragadee Formation; Davidson 1995). Added to this total extraction is the contribution from unlicensed and others (i.e. wetlands, aquifer and cave systems, terrestrial unmeasured residential bores; about one third of all vegetation and river base flow systems). homes on the Swan Coastal Plain own such an appliance Table 1 shows the consequences for each of these types (Government of Western Australia 2007). In sum, these in response to declining groundwater levels and types of extractions, plus declining rainfall and other decreasing rainfall. For instance, perennial streams in the extraneous factors, have resulted in groundwater lower reaches of the Blackwood River are fed by fresh declines of up to 6m on the Mound, and more significant groundwater baseflows (see Anon 2006). These declines in deeper aquifers (Government of Western tributaries are now the refuge for a number of species of Australia 2007). A recent report on the management of freshwater fish, including Balston's Pygmy perch, the Gnangara Mound by the State's Environmental Nannatherina balstoni, which is listed as vulnerable Protection Authority (EPA) gives an estimated loss of 500 under the EPBC Act 1999 (Beatty et al. 2006; Morgan et GL of water over the last 25 years (Anon 2007). al. 1998). Such tributaries also significantly freshen the Elsewhere the EPA pointed out that over the last 25 years more-saline waters in the river, which have derived from existing surface storages had been by down-rated by the cleared middle and upper parts of the catchment. The two-thirds of their capacity, and groundwater allocations groundwater discharge therefore supports not just have increased over the same time to meet demand, populations of freshwater fish in the tributaries, but also including over the past decade. They regarded this flora and fauna in the lower parts of the river dependent situation as not sustainable. upon less saline waters during baseflow conditions (see
5 Journal of the Royal Society of Western Australia, 91(1), March 2008
Table 1 Broad classes of Hydrology - Biology linkages for inland aquatic and terrestrial species, communities and/or ecosystems in coastal margins of the SWAFR (note does not include marine or estuarine systems where groundwater discharge and surface runoff are important). Examples of linkages are given, along with a prognosis under scenarios of declining groundwater levels, decreasing extent and duration of inundation of surface waters, and decreasing rainfall, and examples from the literature where such effects have been detected or predicted.
Hydrology - Biology linkages Examples from SWAFR Prognosis under scenarios of declining groundwater levels and decreasing rainfall
Requirement for seasonal Kwongan vegetation communities Reduced vigour as a result of lower water availability in soil moisture Non-phreatophytic terrestrial summer (Zencich et al 2002), decreased rates of surface soil vegetation carbon and nutrient cycling. Potentially reduced seed set Fringing wetland vegetation and shift in population distribution (Groom et. al 2001) and persistence and community composition (Pettit et al. 2001).
Requirement for seasonally Burrowing or sheltering fauna (ie. Survival provided moisture levels are sustained, however moist habitat for aestivation/ Salamanderfish Lepidogalaxias if seasonal inundation fails, less frequent emergence and drought avoidance salamandmides, see Berra & Allen probably reduced reproduction, may result. (1989) and associated species)
Requirement for a seasonal or Ephemeral wetland systems (i.e. Inundation less frequent (i.e. inundation once every 5 years intermittent surface saturation Sedgelands in Holocene dune to once every ten years) or seasonality of inundation swales of the southern Swan changes (decreasing winter-spring inundation and possible Coastal Plain; Semenuik 2007). incidence of summer inundation). Riparian habitats in riverine Decreased areal extent and duration of inundation can ecosystems, and littoral habitats result in reduced frequency of plant recruitment events of Swan Coastal Plain wetlands. (Pettit & Froend 2001), and reduced richness of wetland invertebrates (J Davis, unpublished data).
Terrestrial requirement for Phreatophytic vegetation (ie. Acute drawdown and low recharge can result in loss of access to groundwater table Banksia woodlands); and the adult individuals of overstorey and understorey species fauna dependent on this (Groom et al 2000) or local extinction of susceptible species vegetation (see Bradshaw et al. (Froend and Drake, 2006). Less severe circumstances can 1999, 2007 for the example of result in reduced vigour of adults and a shift in the the honey possum). distributions of established juveniles (Groom et al. 2001).
Requirement for groundwater 1. Balston's pygmy perch, other 1. Contraction of overall range to habitats of suitability (see discharge to maintain a freshwater dependent riverine Morrissy 1978; Nickoll & Horwitz 1999 for the example of particular quality of surface fauna of tributaries where rivers marron Cherax cainii). water. This quality can be have increased levels of salt 2. Loss of cold stenotherms (mostly Gondwanan stream and influenced by physical (ie. concentrations and lower levels of river fauna; Bunn & Davies 1990; Rutherford et al. 2004). surface water temperatures) dissolved oxygen in summer pools. or chemical (ie. salinity) 2. Most stream and river fauna are characteristics. cold stenotherms and intolerant to elevated water temperature.
Requirement for permanent Permanent lakes, peatlands (ie. Change from permanent to temporary stream systems (ie. surface or subsurface waterbirds requiring drought refuge; loss of Gondwanan fauna; Burnt & Davies 1990). saturation plants wedded to saturated organic Sediments exposed to more frequent drying, potentially rich sediments; potential acid displacing biota. Most severe will be drying heating and sulphate soils). Permanent streams- cracking of sediments that have never been so, changing most Gondwanan fauna has annual sediment structure (Horwitz et al. 1999; Semeniuk & life histories (e.g. dragonflies) Semeniuk 2005) and biogeochemistry; acidification under certain conditions (Sommer & Horwitz 2001).
Requirement for an exchange Hyporheic fauna in river beds and Altered patterns of carbon and nutrient cycling. Reduced between surface/subsurface riparian areas (Boulton et al. 1998) ability to retreat or emerge according to life history flows and groundwater requirement. Potential loss of habitat.
Requirement for saturated Stygofauna Where habitat is fixed at a certain stratigraphic level then hypogean (interstitial) spaces Rootmat communities in caves declines in the saturated zone will strand dependent biota (aquifer) (Jasinska & Knott 2000) resulting in local extinctions (Boulton et al. 2003). Otherwise distributions of short-range endemics may change according to extent of groundwater drawdown (Humphreys 2006).
6 Horwitz et al: Hydrological change stress in a biodiversity hotspot
Pettit et al. 2001 for riparian systems). A decline in water contained in its body (Bradshaw & Bradshaw rainfall (leading to less groundwater recharge) plus 1999). The Honey possum is thus highly dependent on projected declines in groundwater due to local nectar production from Banksia trees (that are themselves extractions, will have the net effect of reducing the extent vulnerable to any fall in shallow water tables; Groom et of freshwater habitat available for Balston's Pygmy perch al. (2000)). in the streams, increasing salinities in the river channel The examples given above, and in Table 1, show the downstream, and presenting the fish with an escalated relevance of direct or indirect reliance of flora and fauna saline barrier to dispersal into the river channel. on water, from surface water and groimdwater to local Within the 38 threatened ecological communities pockets of moisture; they are the higher profile tip of currently listed as endangered or critically endangered hydrology-biology linkages in ecosystems in in Schedules of the Environmental Protection and southwestern Australia. We regard the extraction of Biodiversity Conservation Act (1999) (Department of the groundwater resulting in groundwater declines, and Environment, Water, Heritage and the Arts 2007), 15 climate change, as seriously exacerbating existing threats (39%) are in the SWAFR and 10 of these are found on the such as salinisation, Phytophthora, inappropriate fire Swan Coastal Plain or coastal fringe where urban regimes, and weed proliferation, placing further stress development is occurring. While most are also dependent on already strained ecosystems. on groundwater to some extent (i.e. the Corymbia calophylla - Kingia australis woodlands on heavy soils Becoming a water literate society of the Swan Coastal Plain) some are dearly obligate such A proposal (Anon 2006) to alleviate Perth's perceived as those associated with ironstone outcrops on the Swan water shortage by extracting and transporting 45 GL per Coastal Plain and Scott Coastal Plain (Gibson et al. 2000), annum from an aquifer some 200 km to its south, the tumulus (organic) mound springs of the Swan Coastal Yarragadee Formation, heightened concerns over Plain, aquatic root mat communities in caves north of ecosystems where hydrology-ecology linkages exist, and Perth and in caves on the Leeuwin-Naturaliste Ridge where such impacts have not yet been significant. Models (south of Busselton). predicted that after significant periods of pumping Increased drying associated with groundwater decline groundwater in southern parts of the Swan Coastal Plain will have other biophysical consequences. Periods of and on the Scott Coastal Plain, the water table would inundation will decrease, and periods during which decline where the Yarragadee Formation outcrops sediments will be exposed to drying, will increase. This (Whincup et al. 2005) with concomitant implications for consequence would be of particular concern where biodiversity: in the region of this formation there were sediments would otherwise have been exposed to drying known to be 223 rare and priority flora species and of only very rarely. Shallow potential acid-sulphate soils these a total of 79 taxa had been identified as highly form under anaerobic saturated conditions in the dependent on the maintenance of wetter habitats with presence of iron, sulphur and organic matter. They occur shallow water tables (Anon 2006). Although the State locally but extensively on both the Swan Coastal Plain Government subsequently shelved the proposal on such and the Scott Coastal Plain (Anon. 2003; Degens & environmental grounds, it was replaced with a proposal Wallace-Bell 2006). Where these soils become dry, to build a desalination plant with capacity to produce the acidification is a likely outcome for soils, wetlands and same amount of water. Given the projected increase in groundwaters (Appelyard et al. 2004) posing serious regional population growth, the threat to the aquifer threats to aquatic biodiversity (Sommer & Horwitz 2001). remains unless the dominant ethic of new resource Together, declines of both groundwater and rainfall development is replaced with increased emphasis on the can also have interactive effects with existing threatening systemic relationships between climate change, land use processes. Inappropriate fire regimes, already a concern and local hydrology-biology linkages. (Dixon & Barrett 2003; Hopper 2003; Ho&witz et al. 2003; Carpenter et al. (1992) drew attention to the possibility Yates et al. 2003), may well exacerbate regional drying for global warming to have consequent local and regional and their effects may further alter the frequency and hydrological effects that will have a positive feedback intensity of fires, thus influencing the distribution and effect and enhance global warming. For instance, the abundance of plants and animals. Some, such as the release of greenhouse gases (like methane and carbon endemic trapdoor spiders of the genus Moggridgea, dioxide) from wetland sediments that have dried due to which make cocoon-like tubes on tree trunks and short declining water levels may exacerbate global warming burrows in soil along cuttings and creek banks (Main producing further rainfall declines and lowered water 1999), and the tiny marsupial Honey possum, Tarsipes levels. Southwestern Australia is prone to such rostratus (Bradshaw et al. 2007; Everaardt 2003; Friend & feedbacks: Horwitz ef al. (1999) highlighted the release of Wayne 2003), have been found to be particularly stored carbon from burning dried and exposed organic- susceptible to fire. rich sediments. Other feedbacks will come from any The Honey possum is the only terrestrial vertebrate to realisation of the perceived need to shift to energy depend exclusively on flowers for its pollen and nectar intensive (and greenhouse gas emitting) technologies like diet (Russell & Renfree 1989) and thus is secondarily desalination, or vast interbasin transfers of water, to dependent on water supplies to Banksia that form its produce more water for consumption. Similarly, regional primary food source (Bradshaw et al. 2007). extraction and transport of water will potentially dry Measurements of rates of food intake in free-ranging surface soils at its origin and thereby attract less rainfall. animals show that a 10 g Honey possum consumes These feedback loops are characteristic of systemic almost its own body weight per day in nectar and has a change, and are arguably best dealt with by addressing turnover of water that is over double the total amount of overconsumption of water by domestic, industrial and
7 Journal of the Royal Society of Western Australia, 91(1), March 2008 agricultural sectors while being attentive to the six distal References causes of hydrological change described above. Anon 2003 Acid Sulphate Soils. Western Australian Planning Concerted and coordinated action will be needed by Commission Perth Planning Bulletin 64. natural resource planners and managers if all the threats Anon 2006 South West Yarragadee Water Supply Development: identified to the plant and animal species in Australia's Sustainability Evaluation/Environmental Review & only threatened biodiversity hotspot are not to result in Management Programme Strategen (Glenwood Nominees further losses and extinctions in the coming years Pty Ltd) Perth. (Government of Western Australia 2007). We believe that Anon 2007 Environmental Management of Groundwater sustaining the current and projected populations of Abstraction from the Gnangara Mound. July 2004 - June humans with existing biodiversity in the SVVAFR, in the 2005: Annual Compliance Report Environmental Protection Authority Perth 58. future, will depend on two imperatives: Appleyard S J Davidson W & Commander D P 1999 The effects i. improving our understanding of the way plants of urban development on the utilisation of groundwater and animals survive with the seasonal and inter¬ resources in Perth Western Australia. In: Groundwater in the annual delivery of water in the landscape Urban Environment. Selected City Profiles (ed Chilton). (ecological water requirements); and Balkema Press, Rotterdam, 97-104. Appleyard S J, Wong S, Willis-Jones B, Angeloni J & Watkins R ii. adapting to a different availability of water due to 2004 Groundwater acidification caused by urban rainfall declines, involving different patterns of development in Perth, Western Australia: Source, land- and water-use. distribution, and implications for management. Australian Journal of Soil Research 42:579-585. The aim should be to foster shifts in attitudes to water Atkins K 1998 Threatened flora overview. In: Western and the way we behave with respect to water; carefully Australia's Threatened Flora (eds A Brown, C Thomson-Dans evaluated public information campaigns (as advocated & N Marchant). Department of Conservation and Land by Syme et al. 2000) and the education sector will Management, Perth, 11-14. therefore play a major role in these imperatives Beard J S 1980 A New Phytogeographic Map of Western (understanding and interpreting water as an 'ecological Australia. Research Notes of the West Australian Herbarium literacy' sensu Orr (1992)). Mandatory reductions in the 3:37-58. domestic consumption of water for metropolitan Perth, Beard J S 1999 Southwest Australia. In: Hotspots: Earth's matched by dramatic improvements in irrigation Biologically Richest and Most Endangered Terrestrial efficiencies, will therefore precede further water resource Ecoregions (eds R A Mittermeier, N Meyers, P R Gil & C G developments, and alternatives will be evaluated. For Mittermeier). CEMEX, Mexico, 404-417. instance, Coombes and Lucas (2006) have analysed the Beard J S Chapman AR & Gioia P 2000 Species richness and potential for decentralised options for sustainable water endemism in the Western Australian flora. Journal of Biogeography 27:1257-1268. strategies in urbanised parts of southwestern Australia, including rainwater harvesting, water efficient Beatty S J, McAleer F J & Morgan D L 2006 Fish and crayfish communities of the Blackwood River: Migrations, Ecology appliances and wastewater reuse strategies. In their and Influence of Surface Groundwater. South West analysis of the operation of regional water supply Catchments Council & Department of Water Perth, 185 pp. systems they demonstrated that such strategies could Bell DT 2001 Ecological responses syndromes in the flora of provide significant reduction in regional water demand, southwestern Asutralia: fire resprouters versus reseeders. improvement in regional water security, decreases in Botanical Review 67:417-440. greenhouse gas emissions and economic benefits. Berra T M & Allen GR 1989 Burrowing, emergence, behavior, and functional morphology of the Australian Salamanderfish, Lepidogalaxias salamandroides. Fisheries 14:2-10. Conclusions Bolton G C 1972 A Fine Country to Starve In. University of WA Groundwater declines, reduced surface inundations of Press, Perth, 278 pp. rivers and wetlands and declining rainfall (all predicted Boulton A J, Finlay S, Marmonier P, Stanley E H & Valett H M to continue into the future) are evidence of hydrological 1998 The functional significance of the hyporheic zone in streams and rivers. Annual Review Systematics and Ecology change in the southwestern Australian floristic region. 29: 59-81. This paper constructs eight broad ecological linkages Boulton A J, Humphreys W F & Eberhard S M 2003 Imperilled between hydrology and the biology for the region, and subsurface waters in Australia: biodiversity, threatening with examples demonstrates the extent to which processes and conservation. Aquatic Ecosystem Health & ecosystems will be exposed to increased stresses from the Management 6:41-54. hydrological changes. We recommend that our current Bradby K 1997. Peel - Harvey. 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Western Australia. reviewers, and the assistance of David Blake and Simon Blane in the Australian Mammalogy 29: 25-38. preparation of the manuscript. Brown E M, Burbidge A H, Dell J, Edinger D, Hopper S D &
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Journal of the Royal Society of Western Australia, 91: 13-20, 2008
Origins and nature of vessels in monocotyledons. 10. Boryaceae: xeromorphic xylem structure in a resurrection plant
S Carlquist1, E L Schneider1 & K Kenneally2
'Santa Barbara Botanic Garden, 1212 Mission Canyon Road, Santa Barbara, CA 93105, USA 2School of Earth and Geographical Sciences, Faculty of Natural and Agricultural Sciences, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia.
Manuscript received September 2007; accepted February 2008
Abstract
Boryaceae are unusual in having xylem with an abundance of thick-walled tracheids, combined with relatively few vessel elements with scalariform perforation plates that have very slender bars. These features are relevant to the resurrection plant habit of Borya. The abundance of thick-walled tracheids provide sites for maintenance of water columns during hot dry months, whereas the perforation plates offer minimal impedance to flow during wet periods. This combination is unusual in monocotyledons. Root xylem is like stem xylem in the two species studied. Borya sphaerocephala occurs in the Mediterranean-type climate of south west Western Australia, whereas Borya subulata grows in a tropical summer-monsoon area, the Kimberley plateau of northern Western Australia. Despite differences in climatic regimes of the two species, survival of the dry season appears to be the primary factor in design of the xylem of both, and explains their similarity. Boryaceae are now placed in Asparagales, near Asteliaceae, Blandfordiaceae, Hypoxidaceae, and Orchidaceae. Xylem structure does not parallel the phylogenetic interconnections among these families, in which moist habitats, succulent habits, and other modifications seem to mitigate the effect of ecology in patterning xylem structure.
Keywords: Asteliaceae, Blandfordiaceae, Hypoxidaceae, Orchidaceae, Origin of vessels, scalariform perforation plates, xylem.
Introduction represent stages in xylem evolution generally regarded as primitive (e.g., Cheadle, 1942). The xylem of Borya Labill. (16 described species) and Alania Endl. Orchidaceae (Carlquist and Schneider, 2006) and (2 species) form a family of resurrection plants, Sansevieria (Carlquist and Schneider, 2007) represents Boryaceae, confined to Australia. Earlier recognized primitive conditions that may have persisted by virtue of within Liliaceae (tribe Johnsonieae), or a segregate succulence and other modifications. Boryaceae clearly are family, Anthericaceae, the distinctiveness of Boryaceae not mesomorphic and represent a potential contrast with has been amply confirmed (Chase et al., 1997). Borya those families. plants resemble large mosses in form: they have narrow leaves borne on upright stems. Prop roqts attach them to Various characters of Boryaceae invite comparison to the substrate (sandstone or granite). Little soil is present those of related families. Boryaceae have been considered on the "pavement" areas where they grow, and because to belong to a clade with the order Asparagales that of bright sunlight, the habitats are essentially dry for includes Blandfordiaceae (Blandfordia, 3 species, eastern perhaps half of the year. Borya is commonly considered Australia), Asteliaceae, Hypoxidaceae, and Orchidaceae a resurrection plant, the leaves of which become green (Fay et ail., 2000; Soltis et al., 2000. Cheadle and Kosakai soon after commencement of the rainy season. A number (1971) included Borya nitida Labill. in a monograph of of studies relate to this habit (Gaff, 1981, 1989; Gaff and xylem of Liliaceae. They also list Borya among seven Churchill, 1976; Gaff et al., 1976; Iietherington and species, representing as many genera, of the tribe Smillie, 1982; Hetherington et al., 1982; Kauff et al., 2000; Johnsonieae. Blandfordia, which would no longer be Keighery, 1984). included in that tribe, is illustrated as having long scalariform perforation plates in vessels of roots. Cheadle The groups of monocotyledons the xylem of which and Kosakai (1971) do not illustrate the xylem of Borya, has been studied with SEM in earlier papers of this and only include merged data for Johnsonieae as a whole series, such as Acoraceae (Carlquist and Schneider, 1997) in a table. The xylem of Borya offers excellent material and Araceae (Carlquist and Schneider 1998; Schneider for examination of a question: to what extent are and Carlquist, 1998), represent plants that live in mesic tracheids and vessels in monocotyledons indicators of situations (Carlquist, 1975) and may have had an shared heritage (symplesiomorphies) and to what extent unbroken history in such habitats. They therefore are they templates that reflect adaptation to ecology? What factors hasten or slow changes (or possibly even reverse them) in adaptation to ecological conditions? © Royal Society of Western Australia 2008 Cheadle (1942) has offered a scheme for understanding
13 Journal of the Royal Society of Western Australia, 91(1), March 2008
of evolution of monocotyledon xylem based on Borya sphaerocephala roots (Figs. 1A-1C) morphology. However, the picture he produced Vessel elements have perforation plates that range expresses constraints and pathways. from near-transverse (Fig. 1 A) to markedly oblique (Figs. IB, 1C). In most vessel elements, perforation plates have very slender bars that stretch across the plate. Although Materials and Methods the bars are so tenuous that in macerations they often break (Figs. IB, 1C) or collapse to the margins of the The specimen of Borya, sphaerocephala was collected plates (Fig. 1A), we believe that bars traversing the entire by K. F. Kenneally and K. G. Varnavides on 31 January width of a perforation plate are the rule, and that bars 2007 (PERTH) along a granite apron surrounding the that extend partly across the plate are infrequent. The Darling Scarp, 15 km ENE of Perth (32°01'S 116o01'E), bars taper abruptly from the lateral edges of the adjacent to Bickley Brook Reservoir. The mean rainfall is perforation plate. Perforation plates qualify as ca. 1000 mm in this region, but hot weather (to 37°C) scalariform, with 10-20 bars (possibly some of them renders the shallow sandy soil dry during the summer partial bars). months. B. sphaerocephala flowers during the spring Lateral wall pitting is alternate (Figs. 1A, 1C). Where months (August to October). Plants are often in tufts, less pitting is not evident on lateral walls (Fig. IB, right), the than 10 cm in height. wall may represent an interface with fibers rather than The material of B. subulata was collected by M. D. and with tracheids. R. L. Barrett on 24 January 2007 (PERTH) on a sandstone pavement at Bachstein Creek in the Kimberley Region of Borya sphaerocephala stems (Figs. ID, 2A-2F) northern Western Australia. At this locality, it grows Variation in the width of bars on perforation plates is with Calytrix, Drosera, Eriachne, Eriocaulon, Goodenia, evident: the bars shown in Fig. ID are somewhat thicker Micraira, Triodia, and Utricularia. During the summer than the norm (Fig. 2A), but the range is not very great. rainfall season, the shallow seepage areas where B. Vessel elements mostly have oblique perforation plates subulata grows are covered with 2-3 cm of trickling (Figs. 1D, 2A); a portion near the end of a fibriform vessel water, although the pavement areas are dry during the element is shown in Fig. 2D. winter. Flowering occurs during December to April. The plants are up to 50 cm tall, with stilt roots up to 30 cm Pit membrane remnants may be found at the top and long; longer stilt roots (often carbonaceous) occur on bottom of a given perforation plate, where transitions to plants that grow on bare rock. lateral wall pitting occurs. The pit membrane remnant in Fig. 2B represents an enlargement of the perforation plate Material of both species was collected in the field, then portion shown in Fig. 2A (bottom). Note than an intact preserved in 70% aqueous ethanol. Freehand razorblade pit membrane (wrinkled) occurs at the pit at the bottom longisections of roots were washed with distilled water, of Fig. 2B. The perforation plate portion in Fig. 2E air dried between glass slides, and mounted on represents the top end of the perforation plate shown in aluminium stubs. Macerations were prepared with Fig. 2D; the enlarged portion focuses on a pit membrane Jeffrey's Fluid, washed in distilled water, and air dried remnant in the transitional zone of a perforation plate. on aluminum stubs. In the case of both sections and Pit membrane remnants contain holes of various sizes. macerations, material on aluminium stubs was sputter There are some inconspicuous borders present on the coated with gold and examined with a Hitachi S-2600N perforations shown in Fig. 2E. scanning electron microscope (SEM). Lateral wall pitting of vessel elements from B. sphaerocephala stems is composed entirely of circular bordered pits (Figs. 2C, 2D). These circular bordered pits Results are identical in size and shape to those of the tracheids (Fig. 2F). The pit borders are notably wide compared to Macerations proved more useful than sections for the the pit apertures (Fig. 2C). majority of observations because tracheids much outnumber vessel elements in roots and stems of both Borya subulata roots (Fig, 3) species, and vessel elements are thus insufficiently exposed in sections. Macerations are illustrated in Figs. Roots of B. subulata have vessel elements with oblique 1-3, Fig. 4A-B here, while sections were used to show perforation plates. The tapering of bars is sharp in some pitting of tracheids (Fig. 4C-F). Both vessel elements and plates (Figs. 3A, 3B), suggesting that bars may not extend tracheids have very thick walls that are apparently not across the entire perforation plate. However, in other highly lignified, and thus the oxidative treatment of the plates, one can find portions of very tenuous bars that macerating fluid, followed by washing and drying, have collapsed against the sides of the perforation plates, caused curvature in tracheid shapes, presumably because an inevitable result of the macerating process (Fig. 3C, of the helical distribution of soluble compounds. When left). Determining whether or not slender bars extend such compounds (very likely hemicelluloses and pectins) entirely across a plate is difficult because of the tendency were in part dissolved from walls by the macerating fluid of the plate bars to roll inwards because of the maceration and the tracheids subsequently dried, coiling of tracheids process, obscuring the lateral ends of the bars. resulted. Such coiling was not common in sections. We In the short zones of transition between perforations cannot exclude the possibility that some vessel elements and lateral wall pitting (Fig. 3D), pit membrane remnants in stems might be those of adventitious roots embedded are present. As with the pit membrane remnants in B. in the stem, although we attempted to minimize that sphaerocephala, these remnants are porose. Lateral wall possibility. pitting of vessel elements consists of alternate circular
14 Carlquist et ah Origins and nature of vessels in monocotyledons
Figure 1. Vessel elements of Borya sphaerocephala from macerated roots (A-C) and stems (Fig. D). A. Near-transverse perforation plate; note fine threadlike portions of bars, displaced by the maceration process, at the apparent edges of the perforation plate. B. Highly oblique perforation plate; the margins are incurved, a result of the maceration process. C. Oblique perforation plate; displaced slender bars are present. 4D. Near-vertical perforation plate; borders are faintly visible at the lateral edge of the plate at left. Figs. 1A, IB, scales = 10 pm; Figs. IB, 1C, scales = 20 pm.
15 Journal of the Royal Society of Western Australia, 91(1), March 2008
Figure 2. Portions of vessel elements (A-E) and tracheids (F) from macerations of stems of Borya sphaerocephala. A. Most of a perforation plate, to show attenuated nature of bars. B. Micrograph of the lower portion of the plates shown in A, to show the nature of pit membranes in the bottom two pits (or alternatively, perforations) of the plate. C. Outer surface of a vessel; apertures are oval, similar in shape to the rather wide borders. D. Subterminal perforation plate (below); the vessel element is fibriform. E. Portion of the perforation plate shown in D, to show the nature of the transition region between perforations (below) and lateral wall pitting (above). F. Portions of tracheids; the tracheids have been only partially separated by the macerating process. Figs. 2A, E, F, scales = 10 urn; Fig 2B, scale = 2 pm; Fig. 2C, scale = 5 pm. K B P 6
16 Carlquist et al: Origins and nature of vessels in monocotyledons
Figure 3. Portions of vessel elements from macerations of roots of Borya subulata. A. A perforation plate, margins (especially at right) incurved as a result of the maceration process; bars may be broken from the attenuated tips or else bars may be absent. B. A perforation plate, viewed laterally. C. A portion of a perforation plate, showing that bars are present, but slender portions of them have collapsed against the edges of the perforation plate due to handling. D. A portion of a perforation plate, showing the transition between perforations (below) and pits at the end of the perforation plate; porosities are present in the pit membranes of the three pits (or alternatively, perforations). Figures 3A-3C, scales = 10 pm; Fig. 3D, scale = 5 pm.
17 Journal of the Royal Society of Western Australia, 91(1), March 2008
Figure 4. Portions of vessel elements (A-C) and tracheids (D, E) from macerations (A, B) and sections (C-E) of stems of Borya subulata. A. Perforation plate portion; some bars are intact, others are broken. B. Portion of perforation plate corresponding to the bottom of A, to show the pit membrane remnants in the pits (alternatively, perforations) of the transition area between perforation plate and lateral wall pitting. C. Area below the tip of a vessel element; pits mostly lack pit membranes because of handling; some pits are polygonal in shape. D. Tip of a tracheid, to show the attenuated shape. E. Portions of surfaces of several adjacent tracheids, to show pitting; pit membranes are present. Figs. 4A, 4C-4E, scales = 10 pm; Fig. B, scale = 2 pm.
18 Carlquist et al: Origins and nature of vessels in monocotyledons
bordered pits (Fig. 3A). Tracheids are abundantly present tracheids in orchid xylem promotes potential conductive in roots of B. subulata. safety because of the ability of tracheids to confine embolisms to individual cells. Grasses have minimal Borya subulata stems (Fig, 4) conductive safety, but they have the ability to restore The vessel elements observed in B subulata stems have water columns rapidly if air embolisms have occurred highly oblique perforation plates with 10 or more bars (Stiller et al., 2005). The adaptations of grass xylem and (Fig. 4A). The transitions between lateral wall pitting and of orchid xylem represent modalities quite different from ends of perforation plates (Fig. 4A, bottom; shown those of Borya, a fact that underlines the distinctiveness enlarged in Fig. 4B) features pit membrane remnants. As of Bory'a xylem as representing how the water economy in the pit membrane remnants shown elsewhere in this of a resurrection plant can be achieved. paper (Figs. 2D, 2E, 3S.), porosities of various sizes The slender bars on perforation plates of Borya are penetrate the pit membrane in these areas. Pit membrane distinctive. Based on the morphology of vessels in remnants are confined to transition areas of perforation monocotyledons at large (Cheadle, 1942), one might plates and do not occur in most of the perforations. expect such a vestigial version of the perforation plate Lateral wall pitting of vessels may be so dense that not to be represented so consistently in a genus. Mixtures pits are crowded and polygonal in shape (Fig. 4C) rather of simple and scalariform perforation plates are reported than circular. Tracheids (Figs. 4D, 4E) are much more in some genera and families of monocotyledons abundant than vessel elements in B. subulata stems. (Cheadle, 1942). That suggests that the transition from scalariform to completely simple plates are most commonly abrupt, with preservation of vestigial bars on Conclusions perforation plates a rare condition. Many drawings of perforation plates of monocotyledons have been offered Study of two species of Borya (from a family of 14 or by Cheadle and Kosakai (e.g., Cheadle and Kosakai, more species) may seem a small sample size, but the 1971), but none of their drawings seems to suggest the essential identity of xylem of the two species, despite presence of such tenuous bars on perforation plates. This marked differences between these species with respect to may be an artifact of the drawing method used by habitats and habits, suggests that the patterns described Kosakai, who employed rather coarse lines and may extend throughout the family as a whole. Certainly emphasized simplicity in representation. B. sphaerocephala and B. subulata represent extremes No study of monocotyledon xylem known to us has where habitat and growth form are concerned (see involved determining the relative proportions of Material and Methods). tracheids and vessel elements in xylem of particular Borya is a resurrection plant, a strategy that involves species. In view of the preponderance of tracheids over drying of root, stem, and leaf portions without actual vessel elements in Borya, such analysis becomes highly death of all cells involved (Gaff, 1981, 1989; Hetherington desirable. The assumption, perhaps aided by textbook and Smillie, 1982, Hetherington et al., 1982). The presence drawings of xylem transactions of grass or palm bundles of relatively large quantities of tracheids in xylem of (usually labeled as composed of vessels only) or those of Borya suggests that some water columns remain intact in Yucca stems (usually labeled as composed of tracheids the dry season in tracheid tissue, whereas vessels may only), is that one cell type or the other is present. There is embolize. The pit membranes of tracheids do not permit an interesting parallel in wood of dicotyledons. air bubbles to pass from one tracheid to another (Tyree Vasicentric tracheids occur in various dicotyledon woods: and Zimmermann, 2002). Thick tracheid walls and they may be few in number, or they may (as in Fagacaeae alternate patterns of circular bordered pits are wall and Myrtaceae) be relatively abundant. The entire configurations of maximal strength that would resist the background of a wood may be composed of tracheids. development of high negative pressures that are to be These variations have considerable ecophysiological expected during periods of heat and drought (Carlquist, significance, particularly where xeric habitats are 1975). Vessel elements in Borya, because of their near¬ concerned (Carlquist, 1985). The xylem of Borya, simple perforation plates, would convey water rapidly to therefore, opens a fascinating new avenue for new leaves and to flowering portions of the plant during investigation of xylem of monocotyledons. brief wet seasons. One of the traditional objectives in the study of xylem The xylem reported here for Borya is uncommon in anatomy has been the application of findings to the monocotyledons. Grasses, for example, have xylem that classification system. Indeed, the data in surveys of consists of vessels, with few or no tracheids; all of the monocotyledon xylem have been organized in terms of vessels in grasses have simple perforation plates familial units. Now that Boryaceae is recognized as a (Metcalfe, 1960). The xylem of roots of Orchidaceae family (Chase et al., 1997) belonging to Asparagales, the consists either of vessels (with scalariform perforations most closely related families are considered to be plates) in roots, or else tracheids that have some Asteliaceae, Blandfordiaceae, Hypoxidaceae, and tendencies towards being vessel-like. The stems and Orchidaceae (Fay et al., 2000; Soltis et al., 2000). Indeed, inflorescence axes of orchids have tracheids with some these four families plus Boryaceae might be considered a tendencies toward being vessel-like (Carlquist and suborder of Asparagales according to recent phylogenetic Schneider, 2006). Orchid xylem represents a constructions. The xylem of the four families other than conformation suited for relatively slow rates of water Boryaceae contains vessels with very primitive conduction, a condition related to the succulence, thick perforation plates in roots, and tracheids only in stems cuticles, and other factors likely to promote slow (Cheadle and Kosakai, 1971; Carlquist and Schneider, translocation. In addition, however, the presence of unpublished data). Cheadle and Kosakai (1971) actually
19 Journal of the Royal Society of Western Australia, 91(1), March 2008
do not give illustrations for Borya xylem, and give data Fay M F, Rudall P J, Sullivan S, Stobart K L, De Bruin A Y, only in a merged tabular form for the tribe Johnsonieae Reves G, Qamaruz-Zaman F, Hong W-P, Joseph J, Hahn W J, (of Liliaceae) as a whole, so that our illustrations and Conran J G & Chase M W 2000 Phylogenetic studies of Asparagales based on four plastid DNA loci. In: Monocots: descriptions are apparently the first. The difference systematics and evolution (eds K L Wilson & D A Morrison), between xylem of Boryaceae and that of the four other CSIRO Publishing, Collingwood, Victoria, Australia, 360- families with respect to xylem is conspicuous. This 371. difference corresponds to the unusual nature of Gaff D F 1981 The biology of resurrection plants. In: The biology Boryaceae as a resurrection plant, and should not be of Australian plants (eds J S Pate & A J McComb). University interpreted as evidence of a lack of relationship. DNA- of Western Australia Press, Nedlands, Australia, 114-146. based phylogenies have supplanted comparative Gaff D F 1989 Responses of desiccation tolerant 'resurrection' anatomy as a prime source of evidence for natural plants to water stress. In: Structural and environmental relationships. If we analyze dades of vascular plants in responses to environmental stress (eds J H Kreeb, H. Richter the light of DNA-based phylogeny, we find & T M Hinckley). SPB Academic Publishing, The Hague. unappreciated patterns of diversity in anatomical Gaff D F & Churchill D M 1976 Borya nitida Labill.-an features (Carlquist, 2006). These patterns can be traced to Australian species in the Liliaceae with desiccation-tolerant leaves. Australian Journal of Botany 24:209-224.. the operation of ecology in evolution of tracheary elements and other anatomical features that relate to Gaff D F, Zee S-Y & O’Brien T P 1976 The fine structure of dehydrated and reviving leaves of Borya nitida Labill.-a water economy. desiccation-tolerant plant. Australian Journal of Botany 24:225-236. Hetherington S E & Smillie R M 1982 Humidity-sensitive References degreening and regreening of leaves of Borya nitida Labill. as followed by changes in chlorophyll fluorescencel. Carlquist S 1975 Ecological strategies of xylem evolution. Australian Journal of Plant Physiology. 9:587-599. University of California Press, Berkeley. Hetherington S E, Hallam N D & Smillie R M 1982 Carlquist S 1985 Vasicentric tracheids as a drought survival Ultrastructure and compositional changes in chloroplast mechanism in the woody flora of southern California and thylakoids of leaves of Borya nitida during humidity- similar regions; review of vasicentric tracheids. Aliso 11:37- sensitive degreening. Australian Journal of Plant Physiology 68. 9:601-609. Carlquist S 2006 Asteropeia and Physena (Caryophyllales): a Kauff F P, Rudall J & Conran J G 2000 Systematic root anatomy case study in comparative wood anatomy. Brittonia 58 301- of Asparagales and other monocotyledons. Plant Systematics 313. and Evolution 223:139-154. Carlquist S & Schneider E L 1997 Origins and nature of vessels Keighery G J 1984 The Johnsonieae (Liliaceae); biology and in monocotyledons. 1. Acorus. International Journal of Plant classification. Flora 175:103-108. Sciences 158:51—56. Metcalfe C R 1960 Anatomy of the monocotyledons. 1. Carlquist S & Schneider E L 1998 Origins and nature of vessels Gramineae. The Clarendon Press, Oxford. in monocotyledons. 5. Araceae subfamily Colocasioideae. Botanical Journal of the Linnean Society 128: 71-86. Schneider E L & Carlquist S 1998 Origins and nature of vessels in monocotyledons. 4. Araceae subfamily Philodendroideae. Carlquist S & Schneider E L 2006 Origins and nature of vessels Journal of the Torrey Botanical Society 125:253-260. in monocotyledons. 8. Orchidaceae. American Journal of Botany 93:963-971. Soltis D E, Soltis P S, Chase M W, Mort M E, Albach D C, Zanis M, Savolainen V, Hahn W H, Hoot S B, Fay M F, Axtell M, Carlquist S & Schneider E L 2007 Origins and nature of vessels Swensen A M, Prince L M, Kress W L, Nixon K C & Ferris J S in monocotyledons. 9. Sansevieria. South African Journal of 2000 Angiosperm phylogeny inferred from 18S rDNA. rbcL, Botany 73: 196-203. and atpB sequences. Botanical Journal of the Linnean Society Chase M, Rudall K P J & Conran J G 1997 Validation of the 133:381-461. family name Boryaceae. Kew Bulletin 52:416. Stiller V, Sperry J S, & Lafitte R 2005 Embolized conduits of rice Cheadle V 1 1942 The occurrence and types of vessels in the (Oryza sativa, Poaceae) refill despite negative xylem various organs of the plant in Monocotyledoneae. American pressure. American Journal of Botany 92: 1970-1974. Journal of Botany 29:441-450. Tyree M T & Zimmermann M F 2002 Xylem structure and the Cheadle V I & Kosakai H 1971 Vessels in Liliaceae. ascent of sap. Edition 2. Springer-Verlag, Berlin and Phytomorphology 21:320-333. Heidelberg.
20 Journal of the Royal Society of Western Australia, 91: 21-25, 2008
Greater Bilby (Macrotis lagotis) burrows, diggings and scats in the Pilbara
G G Thompson1 & S A Thompson2
’Centre for Ecosystem Management, Edith Cowan University, Joondalup, Western Australia, 6027, Australia " 0 [email protected] 2Coffey Environments, Dilhom House, 2 Bulwer Street, Perth, Western Australia, 6000, Australia El [email protected]
Manuscript received December 2007; accepted February 2008
Abstract
The greater bilby (Macrotis lagotis) is a conservation significant species because of a contraction in its geographic range and is now only found in a few locations in Australian sandy deserts and the Pilbara of Western Australia. We report on the presence of eight burrows and 605 diggings and scratchings in an area of about 9 ha for a small group of greater bilbies in the Pilbara. We describe burrows that were located adjacent to or under rocks, under an old termite mound and in mature spinifex. Most of the activity area, which had been burnt six months earlier, contained diggings and scratchings that we divided into three categories: nose cone diggings, scratchings and circular holes. Numerous scratchings contained the characteristic scats which can be used to identify the presence of greater bilby in an area. This paper provides information on the evidence that can be used to determine the presence of greater bilby in the Pilbara.
Keywords: Fauna Surveys, Conservation Significant Species, Pilbara, Dalgyte
Introduction It is our experience that the diggings, scratchings and burrows of greater bilbies are occasionally being Greater bilbies (Macrotis lagotis) are listed as confused with that of other species. As this species is vulnerable under the Environment Protection and listed as being of conservation significant with both the Biodiversity Conservation Act (1999) and as a Schedule 1 state and commonwealth governments, a false positive species under the Western Australian Wildlife presence can cause delays or unnecessary costs for Conservation Act (1950). If greater bilbies are potentially developers. In contrast with the habitat utilised by Bilbies present in an area designated for development, then the in the Warburton Ranges, this paper describes one government regulators would expect the developer to aggregation of greater bilby burrows and evidence of determine whether they were present. Given that they foraging in a burnt spinifex meadow on red sandy soil are nocturnal and unlikely to be seen during daytime that may assist others in identifying the presence of searches, evidence from burrows, diggings and scats is greater bilbies during fauna surveys in the Pilbara. particularly useful. Greater bilbies typically use two to three burrows each night (Lavery & Kirkpatrick 1997) and live in small Methods groups of two to four individuals. They dig a burrow A single burrow with four entrances was initially system that may be up to 3 m long and 1.8 m deep located (21° 25’S, 118° 55'E) adjacent to an existing gravel (Johnson 2000). They feed on insects, seeds, bulbs, fruit track that was to be used to haul equipment and and fungi (Smyth & Philpott 1968; Southgate & Carthew materials to a Fortescue Metals Group construction site 2006) and often leave a characteristic excavation of up to in the Pilbara of Western Australia, hr late May - early 10 cm deep with the soil scattered in all directions or a June of 2007 we searched the area on foot to about 100 m 'pot-hole' in the ground (James & Eldridge 2007). Female beyond all located diggings and scratchings. The location greater bilbies reintroduced to a protective environment of all burrows, scratchings and diggings were recorded had a home range of 0.2 km2 and males 3.2 km3 (Moseby with a GPS. The size of each scratching, circular hole and & O'Donnell 2003). Males can grow to 2.5 kg, but females nose cone digging was estimated and recorded. are much smaller with a maximum weight of around 1.1 kg (Johnson 2000). Smyth & Philpott (1968) provided a description of Results burrows, diggings and scratchings of the greater bilby near the Warburton Ranges in a variety of habitat types, A gravel track runs north-south adjacent to the four but most of the burrows were located in mulga-tussock entrance burrow that was initially located (Figure 1). East scrub and mulga-spinifex scrub. There are no data on the of this track the area was mostly vegetated with mature surface signs of greater bilbies for the Pilbara region of spinifex (Figure 2A). West of the gravel track most of the Western Australia. area had been burnt in November 2006 (Figure 2B). There was a small patch of unburnt spinifex (*= 60 m by 50 m) west of the gravel track. A total of eight burrows and 605 diggings and scratchings were identified in an area of © Royal Society of Western Australia 2008 about 9 ha (Figure 1). Two of these burrows were central
21 Figure 1.Distributionofburrows, diggingsandscratchingsofMacrotislagotis
Metres Journal oftheRoyalSocietyWesternAustralia,91(1),March2008 22 Metres Thompson & Thompson: Greater Bilby in the Pilbara
Figure 2. Greater bilby burrow entrances, diggings and scratchings. A - mature spinifex area, B - burnt area, C - nose cone digging, D - scratchings, E - circular hole, F - scats, G - burrow under rocks, H - burrow under rocks, I - burrow under rocks, ] - burrow in spinifex, K - burrow in the open, L - burrow under a termite mound, M - scratchings, N - burrow under rocks, O - scratchings.
23 Journal of the Royal Society of Western Australia, 91(1), March 2008
to the diggings and scratchings, the other six were railway line corridor in July 2006, in an area about 7 km towards the periphery. All burrows showed signs of to the north of this site, a number of greater bilby being recently active. Burrows were judged to have been burrows were located and multiple scratchings and recently active if there was fresh soil around the entrance diggings were found in the vicinity of these burrows in a and footprints in the soil. Some of the scratchings and similar pattern to that described above. This area was diggings had been present for sometime as leaf litter had densely vegetated with mature spinifex and scattered accumulated in the depressions and there were spider trees when initially searched. This area was burnt during webs in a few of the deeper circular holes. November 2006, and when it was searched again in Two burrows were located adjacent to or under rocks December 2006 there was no longer any evidence of (Figure 2G, H, I and N), and one was located under an greater bilbies in the area. We presumed that they had old termite mound (Figure 2L). The burrow with four moved due to a lack of vegetation or an inadequate food entrances was located in mature spinifex (Figure 2J) and supply. the remainder were in the open burnt area (Figure 2K). Burrow entrances were about 300 mm high and 250 mm wide. One burrow with four entrances was within a Discussion couple of metres of the gravel track on the eastern side and one burrow with two entrances under rocks was Greater bilby activity reported here is for a burnt area within a couple of metres of the western edge of the track and we could find no reports of greater bilbies remaining (Figure 2G and I). The piles of soil at the entrance to in burnt areas, however, this may simply reflect the burrows adjacent to the rocks and in the spinifex were paucity of information on their existing habitats. It was about 250 mm high, but those in the open were much unknown how many greater bilbies foraged in this area, smaller. The burrows located were similar to those or how long these burrows had been utilised. However, described by Smyth & Philpott (1968), being slightly oval it was apparent that the foraging area was confined to with a flattened based (Figure 2j and H). The mound of about 9 ha around the burrows. Most of the burrows, soil at the entrance was semi-circular and spread up to diggings and scatchings were located in the area that 1.5 m from the burrow entrance (Figure 2K and N). had been burnt. Tracks were found on the burrow entrances, but they Scratchings and diggings that we report here are were not evident in the surrounding area, probably similar to those described by Smyth & Philpott (1968), because the soil was too hard to make an imprint that being either circular holes or more irregular shapes with would last. lots of shallow diggings into the topsoil only. Smyth & Diggings and scratchings could be grouped into three Philpott (1968) also reported diggings were never more categories: nose cone diggings, scratchings and circular than 180 m from a burrow which again is the same for holes. Nose cone diggings were mostly 'V' shaped with our case study (Figure 1). the deepest section at the apex (Figure 2C). Nose cone It is possible to confuse the diggings and burrows of diggings were mostly 100-150 mm deep. Scratchings other animals with those of bilbies. For example, goanna were mostly shallow (to a depth of about 100 mm) and (Varanus panoptes and V. gouldii) and rabbit diggings, scattered over an area of between 200 by 200 mm to 1200 scratchings and sometimes their burrows can be confused by 500 mm, but mostly 300 by 300 mm (Figure 2D, M with those of greater bilbies. A feature of many goanna and O). Sometimes, a scratching contained one or more diggings deeper than 50 mm, is that they are often nose cone diggings. Circular holes were mostly about 200 adjacent to a vertical spider burrow and the digging mm in diameter and ranged in depth from 200 to 400 slopes into the spider burrow, and can therefore easily be mm, but were mostly about 250 mm (Figure 2E). Circular distinguished from greater bilby holes. However, holes were mostly by themselves, but occasionally in weathered goanna diggings can be confused with those twos and threes. The density of scratchings and diggings of greater bilbies. Greater bilby scratchings and diggings in the areas vegetated with mature spinifex was much are often concentrated around burrows, whereas goanna less than in the burnt area, and there was a higher diggings are much less dense and are not always located density of diggings and scratchings in the centre of the in close proximity to a burrow. Smyth & Philpott (1968) activity area. The ratio of scratchingsmose cone suggested that the opening to greater bilby pits were diggingsrcircular holes was very close to 2:2:1. generally wider (140 mm) than those of goannas (90 Forty three sets of scats were found, mostly on mm). The circular pits dug by greater bilbies at the site scratchings [greater bilby scats are distinctive; being we examined were a little wider (« 200 mm), and pellet shaped and varied in diameter (8 - 13 mm) and although we have not measured those of goannas, our between 10 and 30 mm in length (Figure 2F)J. A set of observations are that they vary appreciably and the scats consisted of 3 to 8 pellets of about the same size and width is influenced by the depth of the digging and age, based on their colour and state of decomposition. probably the size of the goanna that dug the hole. However, it was likely that we missed many of the scats, Greater bilby burrow entrances are generally much as they could have been covered by soil displaced during larger than those of goannas and most other mammals greater bilby digging activity. As scats fade with time, it that use holes in the Pilbara. The burrowing bettong was evident that some scats had been placed on some (Bettongia lesueur), which once lived in the Pilbara, but scratchings on a number of occasions. is now thought to be extinct in the region on the We revisited this site again in September 2007 and the mainland, is the only other mammal with a similar sized burrows were still active, indicating that greater bilbies burrow. Greater bilby burrows are also generally higher can remain in areas with limited vegetation cover. than they are wide, which distinguishes them from During the grid search of the Fortescue Metals Group goanna burrows, which are typically a flattened 'D'
24 Thompson & Thompson: Greater Bilby in the Pilbara
shape with a horizontal base. However, goannas may Acknowledgements: This project was undertaken with the support of the also enter active and inactive greater bilby burrows, and Fortescue Metals Group for which we are very appreciative. in these circumstances their tracks may be obvious around the entrance. References Rabbit (Oryctolagus cuniculus) burrows and warrens can be similar, but smaller, than those of greater bilbies, James A I & Eldridge D J 2007 Reintroduction of fossorial native but their scats (ie circular pellets, ~ 6mm dia, with a lot mammals and potential impacts on ecosystem processes in an Australian landscape. Biological Conservation 138:351- deposited in close proximity) and diggings are very 359. different and difficult to confuse with those of greater Johnson K A 2000 Bilby Macrotis lagotis. In: The Mammals of bilby scratchings which are generally deeper and wider Australia (ed R Strahan). Reed, Chatswood, N.S.W, 186-188. than those of rabbits. Lavery H T & Kirkpatrick T H 1997 Field management of the The presence of greater bilby scats on some of the Bilby Macrotis lagotis in an area of south-western scratchings and diggings is useful for determining Queensland. Biological Conservation 79:271-281. whether the scratchings and diggings belong to greater Moseby K E & O'Donnell E 2003 Reintroduction of the Greater bilbies or another reptile or mammal. Southgate (2005) Bilby, Macrotis lagotis (Reid) (Marsupialia: Thylacomyidae), to Northern South Australia: Survival, Ecology and Notes on demonstrated the relationship between greater bilby scat Reintroduction Protocols. Wildlife Research 30:15-27. diameter and animal size, with heavier animals Smyth D R & Philpott C M 1968 Field notes on rabbit producing larger scats. As the scats we found varied in bandicoots, Macrotis lagotis Reid (Marsupialia), from central size, for example, those shown in Figure 2F ranged from Western Australia. Transactions of the Royal Society of South 8.5 - 12.5 mm, this suggests that different sized greater Australia 92:3-17. bilbies were utilising this area. Southgate R 2005 Age classes of the greater bilby (Macrotis In summary, the presence of greater bilbies can be lagotis) based on track and faecal pellet size. Wildlife Research 32:625-630. determined by an abundance of their characteristic diggings in a confined area, the near circular diggings Southgate R & Carthew S M 2006 Diet of the bilby (Macrotis lagotis) in relation to substrate, fire and rainfall that are about 200 mm in diameter and ranged in depth characteristics in the Tanami Desert. Wildlife Research from 200 to 400 mm, and burrows within the diggings 33:507-519. that have an entrance that is about 300 mm high and 250 mm wide.
25 ' Journal of the Royal Society of Western Australia, 91: 27-30, 2008
Age estimates of Sporolithon durum (Corallinales, Rhodophyta) from Rottnest Island, Western Australia, based on radiocarbon-dating methods
Nisse A Goldberg1-2 & John N Heine3-2
1 School of Plant Biology, University of Western Australia, Crawley, WA 6009, Australia 2 Present address: Department of Biology and Marine Science, Jacksonville University, Jacksonville, FL 32211, USA 3CSIRO Marine and Atmospheric Research, Floreat, WA 6014, Australia
Manuscript received December 2007; accepted February 2008
Abstract
Rhodoliths are free-living coralline algae, found along the coastal waters of temperate and tropical Australia, However, species diversity of rhodoliths has not been investigated to the extent of other subtidal macrophytes. Due to their slow growth rates and longevity, rhodoliths can be aged using radiocarbon dating methods. We aimed to identify species diversity and ages of living rhodoliths collected from Rottnest Island, Western Australia. Rhodoliths were collected from beneath the seagrass canopy of Posidonia spp. and Amphibolis antarctica from Nancy Cove in average depths of 2 m. Rhodolith diversity consisted only of Sporolithon durum (Corallinales, Rhodophyta). Radiocarbon dates from three rhodoliths were estimated using Accelerator Mass Spectrometry. Estimated ages were post-1950, regardless of whether the age was estimated from within 5 mm or up to 15 mm from beneath the rhodolith surface. Age estimates contrast with living rhodoliths collected from deeper waters (depths of 38 m) off the southern coast of Western Australia, whose oldest age estimate was from the year AD 1050. We suggest that seagrass meadows may help facilitate the persistence and growth of rhodoliths growing in the clear and shallow waters of Rottnest Island.
Keywords: rhodoliths, Australia, radiocarbon-dating, encrusting coralline algae
Introduction 2006). In addition, rhodoliths have been recorded from four locations along the coast of Rottnest Island Rhodoliths or maerl are found worldwide with (Playford 1988, Kendrick & Brearley 1997, Sim & records from polar and tropical waters (Foster 2001). Townsend 1999). We aimed to collect and identify Rhodoliths are free-living coralline algae growing living rhodoliths growing in the shallow-waters of subtidally in soft sediments. When they blanket the Rottnest Island; estimate the ages of living rhodoliths seafloor, rhodoliths increase habitat complexity by using Accelerator Mass Spectrometry; and compare providing hard substratum for recruitment of sessile species diversity and ages from our collections to organisms and interstitial spaces for motile animals. published records of rhodoliths located in deeper Individual rhodoliths may be represented by a single waters along the southern coast of Western Australia species or an assemblage of species (Foster 2001). Taxa of (Goldberg 2006). rhodoliths found in Australian waters include Lithophyllum, Lithoporella, Lithothamnion, Hydrolithon munitum, Sporolithon, and Melobesiodeae (a Materials and Methods Mesophyllum or Synarthrophyton) (Collins 1988, Sim & Townsend, 1999, Goldberg 2006). Net growth of a Living rhodoliths were collected from Nancy Cove, rhodolith can result in an increase in rhodolith diameter Rottnest Island (32°02'S, 115°28'W) in average depths of 2 and/or increase in frond length. Net growth may be m, in October 2006. Rottnest Island is located off the uninterrupted resulting in long-lived individuals, or be western coast of Western Australia, in the Indian Ocean. interrupted due to periods of sediment burial (Littler et Local marine communities are influenced by the al. 1991, Goldberg 2006). Growth rates can be relatively southerly-flowing Leeuwin Current, oligotrophic slow with a minimum of < 0.01 mm/yr and a maximum conditions, microtidal changes in seawater levels (< 1.5 m of 2.7 mm/yr, depending on the species and physical range), and exposure to onshore wind flow and southern factors such as water depth (Foster 2001). ocean swells (Collins 1988, Playford 1988). The island In Western Australia, living rhodoliths have been was separated from the mainland of Western Australia about 6500 years ago, is part of a chain of limestone recorded in warm and temperate waters: from the islands and reefs, and is about 10.5 km long and 4.5 km western coast in the IToutman Abrolhos Islands and wide (in Playford 1988). Nancy' Cove and Green Island along the Rottnest Shelf (Collins 1988), and along the are situated along the middle of the south-facing coast of southern coast in the Recherche Archipelago (Goldberg the Island, and are protected from the prevailing southerly winds and swell by offshore limestone reefs (Sim & Townsend 1999). Green Island is about 75 m from © Royal Society of Western Australia 2008 shore.
27 Journal of the Royal Society of Western Australia, 91(1), March 2008
Rhodolith collections were made between Green Sporolithon species (Goldberg 2006). By comparison, only Island and the shores of Nancy Cove. Species S. durum was identified in our study and that of Sim & identifications were made from these rhodolith samples. Townsend (1999) in the shallow waters of Rottnest In addition, we purposely collected 3 large (73, 78, 87 Island. mm in diameter) rhodoliths in order to age each Surprisingly, samples were modern (1950 to present) individual along a cross-section (Fig. 1). Subtidal regardless of whether the section was sampled 0-5 mm vegetative assemblages associated with the rhodolith bed or up to 15 mm in from the surface of the rhodolith consisted of turf algae, encrusting coralline algae, and (Table 1). Resolution was limited by the radiocarbon the seagrasses Posidonia spp. and Amphibolis antarctica. methodology used in this study. Ages could not be Radiocarbon ages of three individual rhodoliths were discerned beyond a timeframe of 1950 (bomb pulse) to estimated using Accelerator Mass Spectrometry (AMS) at present given that percent carbon was greater than 100 the Australian National Tandem Accelerator for Applied pMC (Table 1, pers. comm. V. Levchenko, ANTSO). In Research facility at the Australian Nuclear Science and contrast to the relatively young and larger (>70 mm in Technology Organization (ANTSO). Rhodoliths were diameter) rhodoliths collected from Rottnest Island, age pink and assumed to have been alive at the time of estimates of much smaller rhodoliths (55 mm maximum collection. Dried rhodoliths were first cut in half using a diameter) from Esperance Bay were significantly older water-lubricated diamond saw. Approximately 20 mg with a maximum age estimate of AD1050 as measured was drilled from within 0-5 mm and 5-10 mm of the from 9 mm beneath the rhodolith's surface (Goldberg outer surface along the cut face. Non-algal material was 2006). We recommend using staining methods to carefully avoided. For two rhodoliths, an additional measure growth rates of S. dururm from Rottnest Island sample was collected 10—15 mm from the outer surface. in order to better estimate ages of the shallow-water Samples were prepared for AMS t4C analysis as rhodoliths (Foster 2001). described in Hua et al. (2001): rhodolith samples were In Nancy Cove, rhodoliths were found interspersed converted to carbon dioxide and then to graphite. The within a seagrass meadow (Fig. 2). We suggest that AMS ,4C results were calibrated with the marine seagrasses may facilitate the persistence of rhodoliths calibration curve (Hughen et al. 2004), and corrected for growing in shallow and non-turbid waters. Seagrass 8 13C isotopic fractionation and local oceanic reservoir meadows stabilize sediments and can serve as a baffle to (reservoir correction: 323 ± 86 years; approximate AR attenuate wave energy (Piller & Rasser 1996), thereby value of 25 ± 1 pMC; Gillespie & Polach 1979, V. minimizing the potential for burial of rhodoliths (Steller Levchenko, ANTSO, pers. comm.). & Foster 1995). The ellipsoidal shapes of rhodoliths from Nancy Cove were similar to those collected from a bed in the Red Sea, Egypt with a comparable physical habitat Results and Discussion (depths of 1 to 3 m, exposure to a microtidal climate, protection from wave energy by adjacent reefs, presence The rhodolith bed from Nancy Cove, Rottnest Island, of a seagrass meadow) (Piller & Rasser 1996). Insufficient consisted of rhodoliths interspersed among seagrass water motion may prevent rolling of larger rhodoliths blades in average depths of 2 m (Fig. 2). Individual (lower surface was paler pink compared to the upper fragments in the rubble were approximately 10-20 mm surface) as suggested by Piller & Rasser (1996) (but see in length, and larger individuals ranged in diameter from Steller & Foster 1995). 70 to 220 mm. Kendrick & Brearley (1997) had reported spherical rhodoliths at Nancy Cove with diameters of Despite the year-round presence of a seagrass canopy, 100 mm. In our collections, rhodoliths had nuclei of the leaves may not inhibit recruitment of coralline algae. coralline algae and overall morphology was ellipsoidal to Sim & Townsend (1999) had recorded successful spherical. recruitment of coralline algae under a kelp canopy in Rhodoliths consisted of one species, Sporolithon durum (Foslie) R.A. Townsend & Woelkerling, from our collections. In Australia, Sporolithon durum Table 1 (Sporolithaceae, Corallinales, Rhodophyta) has been Age estimates from three living rhodoliths that were collected reported from Western Australia (Rottnest Island), New from depths of 2 m at Rottnest Island, Western Australia. South Wales (Byron Bay, Coffs Harbour, Jervis Bay, Two¬ fold Bay, Port Stephens, Botany Bay), and in Victoria ANTSO* Rhodolith Distance A(13C) Conventional (Gabo Island) (Woelkerling 1996, Harvey et al. 2002). At code individual (mm) from per 1JC age (month Rottnest Island, Sim & Townsend (1999) identified (maximum rhodolith mil and year rhodoliths of S. durum from depths of 2 to 36 m. In even diameter) surface graphite + la error) deeper waters (< 60 m) at the Rottnest Shelf, rhodoliths OZJ346 1 (78 mm) 0-5 -0.8 1950 to present consisting of the genus Lithothamnion had been recorded OZJ347 1 5-10 -1.7 1950 to present (Collins 1988). OZJ348 2 (73 mm) 0-5 -0.7 1950 to present Species diversity of coralline algae that made up OZJ349 2 5-10 -1.3 1950 to present individual rhodoliths differed between the shallow-water OZJ350 2 10-15 0.9 1950 to present (depths of 2 m) bed sampled in our study and the deeper- OZJ351 3 (87 mm) 0-5 -1.0 1950 to present OZJ352 3 5-10 -2.0 1950 to present water (depths of 35 m) bed sampled in Esperance Bay, OZJ353 3 10-15 -0.8 1950 to present Western Australia (Goldberg 2006). Species richness of rhodoliths growing in Esperance Bay consisted of *ANTSO: Australian Nuclear Science and Technology approximately four taxa, including the possibility of a Organisation
28 Goldberg & Heine: Age estimates of Sporolithon durum, Rottnest Island
Figure 1. Sporolithon durum rhodolith (ANTSO code OZJ351-353 from Table 1).
Figure 2. Rhodoliths in seagrass meadow.
29 Journal of the Royal Society of Western Australia, 91(1), March 2008
Nancy Cove. Instead, seagrass blades may minimize the Goldberg N 2006 Age estimates and description of rhodoliths potential of overexposure to irradiance by S. durum, from Esperance Bay, Western Australia. Journal of the which had also been recorded in depths of 36 m (Sim & Marine Biological Association of the United Kingdom 86:1291-1296. Townsend 1999). Thus, seagrasses may facilitate the existence and persistence of rhodoliths in shallow-waters. Harvey A S, Woelkerling W J & A J K Millar 2002 The Sporolithaceae (Corallinales, Rhodophyta) in south-eastern The persistence of rhodolith beds in Nancy Cove may Australia: taxonomy and 18S rRNA phylogeny. Phycologia rely upon vegetative fragmentation of rhodoliths and 41:207-227. settlement of propagules from S. durum populations (Sim Hua Q, Jacobsen G E, Zoppi U, Lawson E M, Williams A A, & Townsend 1999). Rhodolith rubble (diameters < 10 Smith A M, & McGann M J 2001 Progress in radiocarbon mm) may represent a substratum for spores or gametes target preparation at the ANTARES AMS Centre. to settle and grow. Investigations of rhodolith growth Radiocarbon 43:275-282. rates, reproductive potential, individual survivorship, Hughen K A, Baillie M G L, Bard E, Bayliss A, Beck J W, and fate of fragments may help clarify processes that Bertrand C J H, Blackwell PC, Buck C E, Burr G S, Cutler K B, Damon PE, Edwards R L, Fairbanks R G, Friedrich M, contribute to the persistence of the Nancy Cove rhodolith Guilderson T P, Kromer B, McCormac F G, Manning SW, bed. Bronk Ramsey C, Reimer P J, Reimer RW, Remmele S, In summary, living rhodoliths collected in shallow Southon J R, Stuiver M, Talamo S, Taylor FW, van der Plicht J & Weyhenmeyer C E 2004 Marine04 Marine radiocarbon waters from Rottnest Island were species depauperate age calibration, 26-0 ka BP. Radiocarbon 46:1059-1086. and young compared to deep-water rhodoliths collected Kendrick G A & Brearley A 1997 Influence of Sargassum spp. from the south coast of Western Australia. The rhodoliths attached to rhodoliths on sampling effort and demographic from Rottnest Island were interspersed under a seagrass analysis of Sargassum spp. (Sargassaceae, Phaeophyta) canopy. The seagrass meadow may help minimize the attached to a reef. Botanica Marina 40:517-521. effects of solar inhibition (i.e. possible photodamage) and Littler M M, Littler D S & Hanisak M D 1991 Deep-water wave action (burial and excessive fragmentation). The rhodolith distribution, productivity, and growth history at presence of rhodoliths in the seagrass meadow may also sites of formation and subsequent degradation. Journal of increase local productivity and diversity due to increased Experimental Marine Biology and Ecology 150:163-182. microhabitats for a variety of macroscopic and Piller W E & Rasser M 1996 Rhodolith formation induced by microscopic flora and fauna. reef erosion in the Red Sea, Egypt. Coral Reefs 15:191-1998. Playford P E 1988 Guidebook to the geology of Rottnest Island. Geological Society of Australia, WA Division and the Acknowledgements: We would like to thank the Rottnest Island Geological Survey of Western Australia, Perth. Authority for providing permission to collect, Australian Institute of Sim C B & Townsend R A 1999 An account of common crustose Nuclear Science and Engineering for funding support (grant 06/070), and the Australian Nuclear Science and Technology Organisation for dating coralline algae (Corallinales, Rhodophyta) from macrophyte the rhodoliths. In particular, we would like to thank A. Jenkinson, V. communities at Rottnest Island, Western Australia. In: Levchenko, and G. Jacobsen (ANTSO), and R. Scott (UWA). R. Riosmena- Proceedings of the ninth International Marine Biological Rodriguez confirmed our algal identification. Workshop: The seagrass flora and fauna of Rottnest Island, Western Australia (eds D.I. Walker & F.E. Wells). Western Australian Museum. 1:395-408. References Steller D L & Foster M S 1995 Environmental factors influencing distribution and morphology of rhodoliths in Bahia Collins L B 1988 Sediments and history of the Rottnest Shelf, Concepcion, B.C.S., Mexico. Journal of Experimental Marine southwest Australia: a swell-dominated, non-tropical Biology and Ecology 194:201-212. carbonate margin. Sedimentary Geology 60:15-49. Woelkerling W J 1996 Corallinales. In: The Marine Benthic Flora Foster M S 2001 Rhodoliths: Between rocks and soft places. of Southern Australia (ed H B S Womersley). Australian Journal of Phycology 37:659-667. Biological Resources Study, Canberra, 146-322. Gillespie R & Polach H A 1979 The suitability of marine shells for radiocarbon dating of Australian prehistory. In: Radiocarbon dating: proceedings of the 9th international conference on radiocarbon dating (eds R Berger & H Suess). University of California Press, Los Angeles and La Jolla, 404-
30 Journal of the Royal Society of Western Australia, 91: 31-50, 2008
A new species of soldier crab, Mictyris occidentalis (Crustacea: Decapoda: Brachyura: Mictyridae) from Western Australia, with congener comparisons
J Unno 2
1. School of Natural Sciences, Edith Cowan University, Joondalup, WA 6027 2. V & C Semeniuk Research Group, 21 Glenmere Road, Warwick, WA 6024
Manuscript received November 2007; accepted February 2008
Abstract
Mictyris occidentalis, a new species of mictyrid crab (commonly known as soldier crab), is described from Western Australia. Diagnostic features include the macroscopically granular carapace; prominent granular ridges running posteriorly from the antero-lateral spines; front with broadly curved lateral lobes; rounded and projecting shape of the carapace posterior border and distinctly curved and setose dactyls on the 4th pair of walking legs. The new species combines characteristics of congeners M. longicarpus Latreille, 1806 and M. brevidactylus Stimpson, 1858 with which it is compared biometrically. M. occidentalis is the only Mictyris species extant from Shark Bay to Broome in Western Australia and is likely to be endemic. The taxonomic history of Mictyris species is summarised, and recognition of M. brevidactylus as a separate Asian species/ species complex and M. longicarpus as endemic to the eastern coast of Australia is supported. A key to the described species of Mictyris is provided.
Keywords: Intertidal crab, Mictyridae, Mictyris occidentalis, new species, soldier crab, taxonomy, endemic. Western Australia.
Introduction 1985, 2002). Habitat, behaviour and distribution of M. occidentalis also are described briefly. Soldier crabs (Mictyris spp.) are well-known The new species is morphologically and biometrically inhabitants of tidal flats and beaches in the central Indo- compared with two other Mictyris species, west Pacific region, highly visible in large roving M. longicarpus and M. brevidactylus. All three species of "armies" of several hundred small, round blue crabs Mictyris examined in this study were previously emerging during low tide. They are also distinctive in their unique ability among the Brachyura to walk (or assigned by McNeill (1926) to either M. longicarpus, including Mictyris collected from the Western Australian run) forwards, the presentation of the cheliped carpus in a vertical plane (similar to a praying mantis), and their coast (i.c., the M. occidentalis of this study), or in the case of M. brevidactylus Stimpson, 1858, to a variety of habit of rapidly burrowing into the substrate in a M longicarpus (viz. M. longicarpus var. brevidactylus) corkscrew motion. Besides the Mictyris species, the restricted to South-east Asia. To determine clearly common name "soldier crab" has also been applied to other crustaceans such as hermit crabs (the European whether there is one species with varieties (as McNeill suggested), or there are three separate species in this Parurus bernhardus and the Caribbean Coenobita congener group, it was necessary to undertake detailed clypteatus), the ocypode crab Dotilla myctiroides from comparisons between the three forms. South-east Asia and, occasionally, the Christmas Island Red Crab, Gecarcoidea natalis, however, the term The paper also provides a key to help differentiate "soldier crab" is most commonly applied to species of between the five species of Mictyris described to date. Mictyris. A brief review of the taxonomic history of Mictyris and its inherent problems is provided as background for Taxonomic history the description of the new species. While the Family Mictyridae Dana, 1851 has its own This paper describes Mictyris occidentalis, a new convoluted history (e.g., Crane 1975 refers species from the monotypic Family Mictyridae Dana, "Myctyris"[sic] to the Ocypodidae; Jones 2004 similarly 1851, consistent in morphological characteristics across a referred Mictyris that occurred in Withnell Bay, Western distribution recorded thus far from Monkey Mia, Shark Australia to the Ocypodidae), this paper is concerned not Bay to One Armed Point, near Cape Leveque on the with the history and status of the Family but only with mainland coast of Western Australia (Fig. 1). As such, it Mictyris itself as a context to the identification of a new is a contribution towards clarifying a part of the complex species. However, this section on taxonomic history does of species referred to Mictyris longicarpus (Davie 1982, not attempt an exhaustive treatment of synonymies on species of Mictyris, which can be found in the literature, but rather focuses on the problems in taxonomy resulting © Royal Society of Western Australia 2008 in M. occidentalis being originally assigned to
31 Journal of the Royal Society of Western Australia, 91(1), March 2008
differences have become apparent. Besides imprecise 115 EI 125 EI distributions, other problems included limited access to N literature in foreign languages, particularly early references, and the creation of synonymies. The misspelling of species names in the literature also added to the taxonomic confusion of some groups, including the Mictyridae (Alcock 1900). The genus Mictyris with its holotype Mictyris longicarpus and the nebulous type locality of "Oceano Indiae orientalis", was originally described by Latreille in 1806 from a collection in the Paris Museum. Latreille was not the collector, hence the vagueness of the type locality, a problem which still has repercussions for researchers today. To Latreille's credit, efforts were made to clarify the type location and later the locality was amended to " l'Ocean australasien" (Latreille 1829). The species Mictyris longicarpus was referred to by subsequent authors under numerous misspellings, e.g., "Myelitis longicarpis" (Milne Edwards 1837), before being corrected to the original name by Alcock (1900) who also gave a more detailed description of the genus. Dana (1851) elevated the genus to family status, a move supported by Kemp (1919), McNeill (1926), and most of the recent brachyuran catalogues (e.g., Davie 2002). A second species of Mictyris in Australia, M. platycheles, was described by Milne Edwards (1852) from Tort Western' (Western Port Bay), Victoria with a distribution from Moreton Bay, Queensland to the north coast of sampling site for holotype, allotype; Tasmania. The nomen nudum Myelitis subverrucatus • biometrics for 5 individuals;carapace was given to a specimen from Tasmania by White in length for several hundred individuals 1847 but since has been assigned to the synonymy of M. platycheles along with the provisional name Myelitis prostoma suggested by Stimpson in 1907 (McNeill, 1926). sampling sites for biometrics for De Haan (1835) published the name Mictyris deflexifrons o 5 individuals;carapace length for (recognised now as a nomen nudum) for specimens from several hundred individuals Japan (which are now assigned to M. brevidactylus, see below) sampling sites for measurement o of carapace length for several The next important stage in the history of Mictyris hundred individuals taxonomy was the work of Stimpson (1858) who described very briefly, but did not illustrate, Mictyris brevidactylus from the southern coast of China and the Figure 1. Map of Western Australia showing location and types islands of "Loo Choo" (Ryukyu Islands, Japan), and of sampling sites for Mictyris occidentalis. definitively established that the South-east Asian Mictyris did not belong to M. longicarpus. Unfortunately, M. longicarpus, and in M. brevidactylus from South-east the types of M. brevidactylus were lost in a fire at the Asia being confused with M. longicarpus even today, Chicago Academy of Sciences in 1871 and William more than 200 years since the holotype was first Stimpson died in 1872 without publishing a full described. description and figure of the new species. In 1907, Dr Mar)' J. Rathbun, who, as assistant curator of the United The taxonomic history of Mictyris is similar to that of States National Museum, was familiar with Stimpson's a multitude of taxa which were originally described in concept of M. brevidactylus, published a posthumous the early 19,h Century from collections made on European paper by Stimpson which described and contained exploratory voyages to the Indian-Pacific region. Early figures of M. brevidactylus, and gave its distribution diagnoses were often very brief due to the large number from Hong Kong, South China Sea, to the Ryukyu of organisms collected and there was a tendency to place Islands, southern Japan (Stimpson 1907). However, this superficially similar specimens into existing groups paper (Stimpson 1907) did not eliminate the confusion rather than create new taxa. Frequently, type locations over the misuse of the name "M. longicarpus" in South¬ from early voyages were regional (e.g., "east Indian east Asia and Japan. Adding to the taxonomic confusion, Ocean ), rather than site-specific, or sometimes were De Man (1887) and Tesch (1918) regarded incorrectly recorded, and often specimens were described M. brevidactylus Stimpson to be synonymous with at a later date, not by the collector but by another worker, M. longicarpus, although McNeill (1926) argued that usually a museum taxonomist. Consequently, over time, these workers were not in possession of specimens of many taxa have undergone numerous revisions and have true M. longicarpus from eastern Australia for proper been re-assigned by various authors, as species comparison, unlike Stimpson who had collected material
32 Unno: New species of soldier crab
from Port Jackson and Botany Bay, in New South Wales "M. longicarpus" from the Ryuku Islands with (NSW). M. longicarpus from New South Wales and regarded them as "specifically distinct". Currently, some workers McNeill (1926), in a significant revision of the accept Mictyris brevidactylus as a synonym or just a Mictyridae, recognised three species and one variety of variety of M longicarpus (e.g., Kawaguchi 2002), while soldier crab in the lndo-Pacific and Australian regions. others, notably most Australian, Japanese, Chinese and These were: Mictyris longicarpus Latreille, 1806; Mictyris South-east Asian researchers (e.g., Shih & Liao, 1998; platycheles H. Milne Edwards, 1852; Mictyris Takeda, 2005) accept the designation of Stimpson (1858, livingstonei McNeill, 1926 (type locality at Trial Bay with 1907). a distribution from Cooktown, Queensland (Qld.) to Trial Bay, northern NSW), and Mictyris longicarpus var. Davie (1982, 1985, 2002) suggested that Mictyris in brevidactylus (Stimpson). McNeill did not give a Australia is a "complex" of five species: three from rationale for changing the status of M. brevidactylus, eastern Australia have been described (M. longicarpus although he did suggest that future workers would find Latreille, 1806, M. platycheles H Milne Edwards, 1852 local "definite racial forms" within the distributional area and M. livingstonei McNeill, 1926); one from the of the variety. In addition, with regard to the identity of Northern Territory remains undescribed; and the fifth true M. longicarpus, McNeill (1926) provided a detailed from Western Australia is the subject of this paper. argument for the type location of Latreille's This present study strongly supports recognition of M. longicarpus as Port Jackson (NSW), on the east coast full species status for M. brevidactylus Stimpson, 1858 of Australia. McNeill set the distribution of and, following Davie (2002), M. longicarpus (sensu M. longicarpus (in Australia) as being from the lower stricto) as being restricted to eastern Australia. Other NSW coast northwards, across the northern part of records of M. longicarpus from elsewhere in Australia Australia and along the coast of Western Australia to and the Indo-west Pacific Ocean are either Perth in southwestern Australia, and stated that soldier misidentifications or are of undescribed species. crabs from the Philippine Islands and northwards to the China Sea should be referred to M. longicarpus var. brevidactylus. The record from Perth (Miers 1884), on Materials, methods and terminology which McNeill (1926) rests the southwestern-most occurrence of M. longicarpus in Western Australia, is Materials examined: unlikely to be correct as there are no other records of Mictyris further south than Shark Bay along the Western Abbreviations: Australian mainland coast. [1 have not found Mictyris 6 = male, $ = female, J = juvenile, ovig = ovigerous, south of Shark Bay on the Western Australian mainland AM = Australian Museum, WAM = Western Australian coast in extensive surveys of appropriate sandy tidal flat Museum, VCSRG = V & C Semeniuk Research Group, habitats in the estuaries from Kalbarri to Perth between max = maximum, CL = carapace length 1997 and 2007], Of particular relevance to this paper was McNeill's Material for comparison and biometric study. treatment of a series of specimens from the general Mictyris occidentalis: 4 6 3 (CL 12.90-12.73 mm) locality "North West Australia" including Cygnet Bay WAM C39422, King Bay, Dampier Archipelago, WA, north of Broome in Western Australia. Despite observing 2004; 5 66 (CL 14.96-13.36 mm) WAM C39423, Broome that the "greatest divergence from the typical form" WA, 2004; 5 66 (CL 12.95-10.52 mm) WAM C39424 occurred in this series, McNeill (1926) conservatively Monkey Mia, Shark Bay, WA, 2004 assigned the Western Australian populations of soldier crabs to M. longicarpus. Tire characters which McNeill Mictyris brevidactylus: 3 6 6 ( CL 12.01-9.07 mm), 3 listed for "North West Australia" are consistent with 2 9 (CL 11.32-8.47 mm) AM 5750 Subig Bay, north M occidentalis, although McNeill does not mention the Philippine Islands, 1908; 4 6 6 (CL 15.25-11.14 mm) AM unique projection of the carapace posterior border or the P73642 Starfish Bay, New Territories, Hong Kong, 2006 gonopod morphology (as described in this paper). As Mictyris longicarpus: 7 66 (CL, 19.58-11.23), 3 2 2 such, the taxon "M, longicarpus (pro parte)" referred to (CL 15.15-13.12) WAM C13323 Cape Ferguson, in the synonymy of this paper (see below) refers to those Townsville, Queensland, Australia, 1982 Mictyris crabs from Western Australian that were assigned to M. longicarpus by McNeill (1926). Material from Australian localities: Notwithstanding the revisions by McNeill (1926), later M. occidentalis in the VCSRG collection: 4 6 6 (CL workers, especially in the Japanese and South-east Asian 12.90-12.73 mm), 2 non ovig 2 9 (CL 10.6-9.2 mm), 1 regions, still held to Latreille's initial broad type locality ovig 9 (CL 8.80 mm), 32 JJ (CL 6.00-1.20 mm) VCSRG (i.e., M. longicarpus occurred everywhere, from Japan to 80804 Coconut Well, WA, 2004; 62 6 <3 (CL 15.10-7.30 Australia). Sakai (1976), for instance, stated that Japanese mm), 3 non ovig 2 9 (CL 10.40-9.20 mm), 92 JJ (CL 4.70- crab fauna contained a single Mictyris species, 1.50 mm) VCSRG 70804, Broome, WA, 2004; 2 2 VCSRG M. longicarpus, and agreed with Tesch (1918) that 110294 Port Smith, WA, 1994; 255 66 (CL 13.90-7.10 M. brevidactylus was synonymous with M longicarpus. mm), 192 non ovig 2 2 (CL 10.50-7.10 mm), 1 ovig 9 At that time, Japanese carcinologists referred Japanese (CL 9.40 mm), 68 JJ (CL 6.50-1.40 mm) VCSRG specimens to M. longicarpus as they had not accessed 120704COSS, Settlers Beach, Cossack, WA, 2004; 2 6 6 McNeill's paper which limited the distribution of true (CL 7.80-7.70 mm), 1 non ovig 2 9 (CL 6.70 mm), 179 JJ M. longicarpus to Australia (Takeda 1978). Takeda (CL 5.80-1.40 mm) VCSRG 140704 Point Sampson, Cape (1978), citing McNeill's 1926 paper, compared Lambert, WA, 2004; 37 66 (CL 12.90-7.00 mm), 5 non
33 Journal of the Royal Society of Western Australia, 91(1), March 2008
ovig $ 9 (CL 10.90-7.00 mm), 5 ovig 9 9 (CL 8.90- M. longicarpus for accurate taxonomic comparison of 7.00 mm), 272 JJ (CL 5.70-1.50 mm) VCSRG 150704 characters. Hearson Cove, Dampier Archipelago, WA, 2004; 152 JJ To determine which soldier crab species were extant (CL 6.40-1.50 mm), VCSRG 120704WB Withnell Bay, in Western Australia, circa 1000 specimens were collected Dampier Archipelago, WA, 2004; 295 6 6 (CL 13.90-7.00 mm), 208 non ovig 2 2 (CL 10.90-7.00 mm), 5 ovig 2 9 from populations in 18 sub-tropical to tropical coastal (CL 9.40-7.00 mm), 527 JJ (CL 6.90-1.00 mm) VCSRG- sites for morphological and biometric studies (Fig. 1 and Table 1, all co-ordinates in WGS84 system). DA, Dampier Archipelago, WA, 2001-2003; 2 66 (CL 8.20-7.50 mm), 5 ovig 2 2 (CL 9.90-7.50 mm), 4 JJ (CL 3.90-3.20 mm) VCSRG 1107040NS Onslow, WA; 42 6 6 (CL 15.80-7.30 mm), 31 non ovig 2 2 (CL 12.50-8.60 mm), 10 ovig 2 2 (CL 9.90-7.50 mm), 467 JJ (CL 5.60-1.20 Table 1 mm) VCSRG 110704CARN Carnarvon, WA; 23 6 6 (CL Broome to Shark Bay regional study collection sites from 15.40-7.00 mm), 32 2 2 (CL 12.20-6.80 mm), 4 JJ (CL localities in WA 6.50-5.50 mm) VCSRG 100704MM Monkey Mia, Shark Bay, WA; 3,995 6 6, 2 2, JJ (CL 13.9-1.0 mm) VCSRG Region Locality/Site Latitude 19802003 King Bay, Dampier Archipelago, WA, 1980- & Longitude 2003. Broome Coconut Well 17° 49’ 16.37" S M. occidentalis in the WAM Collection: 1 6 (CL 122° 12' 44.78" E 13.09 mm) C39414 One Armed Point, NW Kimberley, Riddell Beach WA; 1 6 (CL 14.15 mm) C13294 Gnamagun (Lombardine 17° 59' 05.15" S 122° 11' 28.02" E Mission) WA, 1982; 2 2 9 WAM Cl 2970, 1980 & C16995, 1978, Barrett Creek NW Broome, WA; 6 6, 9 2, JJ (max Broome Town 17° 58' 02.47" S CL 15 mm) Cl 2979, Cl 2940, C12934, C39405 Broome, high tide 122° 14' 16.34" E WA, 1980; <3d, 2 2, JJ (max CL 11.71 mm) C8037, Broome Town 17° 58' 02.47" S C39398, C39399 & C39415 Mundabullangana, near Port mid-tide 122° 14' 16.34”E Hedland, WA, 1960; 6 6 C13009 & C13010 Mko Bay Port Hedland Cape Lambert, WA, 1979; 1 6 (CL 12.14 mm) C39408 Port Hedland 20° 18’ 26.80" S spoil bank Legendre Is. Dampier Archipelago WA 1962; 6 6 C25898, 118° 35' 28.30" E C26685, C29204 Dampier Archipelago, WA, 1998-99; 66 6 Mile Creek 20° 19’ 24.90" S (max CL 15.27 mm). Cl 1871, C23036, C23354, 03001-05, 118° 39' 52.56" E C38788-90, C39390, Ningaloo/Exmouth Gulf region WA, 12 Mile Creek 20° 20' 09.60" S 1974-95; 6 6 (max CL 14.00 mm) C5689, C22231, C39402,’ 118° 40' 20.89" E C39409-10, Carnarvon region, WA, 1939-76; 1 6 (CL 15.53 mm) Faure Is., Shark Bay, WA, 1958; 2 9 6 6 (max Cape Lambert Settlers Beach 20° 40' 03.20" S CL 17.49 mm) C8038, Monkey Mia, Shark Bay, WA, 1960. (near Cossack) 117° 14' 40.24" E M. longicarpus in the WAM Collection: 1 2 (CL Point Samson 20° 37' 47.46" S 13.27 mm) 1 C39392, Sandfly Creek, MacKay, QLD, 1983; 117° 11'51.04" E 1 6 C8039 Townsville, QLD 1961; 1 ovig 2 C13327 Dampier Archipelago Hearson Cove 20° 37' 37.63" S Townsville, QLD, 1982; 1 M. livingstonei in the WAM Collection: 1 J, C39401, Hearson Cove 20° 37' 42.14" S Sandfly Creek, MacKay, QLD, 1983; 1 6 C39416" sand flat 116° 47' 51.41" E Andergrove, QLD; 1 M platycheles in the WAM Collection: 1 6 C39396 King Bay-King Altona Pier, VIC, 1965 20° 38' 0.8.56" S Bay Shoal 116° 45' 25.48" E Mictyris sp. in the WAM Collection: 4 6 6 (CL 15.38- King Bay-KB12 20° 38' 10.38" S 12.01 mm), 2 2 2 (CL 11.54-11.05 mm), 19116 NW of 116° 45' 28.87" E Unwins Island, north Kimberley, WA, 1988 (Mictyris Ashburton River Delta longicarpus is on the Museum label); 1 J C39389 Lee Onslow 21° 38' 52.28" S Point, Darwin, NT; 1 «J (CL 12.00 mm) C39395 West 115° 07' 50.74" E Woody Island Creek Gove, NT Gascoyne River Delta One Tree Road 24° 51' 43.40" S 113° 37' 43.90" E Material from non-Australian localities: Mile Jetty 24° 54' 09.85" S Mictyris brevidactylus in the WAM Collection: 6 6 C39406, C39412, C39413 Hong Kong 113° 40' 01.97" E Mile Jetty West 24° 54' 09.85" S Taxonomic work and morphological descriptions and 113° 40' 01.97" E comparisons in this study were based on: (1) data from Shark Bay several hundred specimens collected and examined Carnarvon 24° 54' 15.1m4" S racecourse 113° 40' 19.18" E “ f fIfc!atitL'dlnnI g^lent in Western Australia (25 S to 16 S); and (2) a particular focus on obtaining Monkey Mia 25° 47' 36.44” S correctly identified specimens of M. brevidactylus and 113° 43' 18.05" E 34 Unno: New species of soldier crab These collection sites ranged from Monkey Mia, Shark Similarly, it was considered important to obtain Bay in the south to Coconut Well (north of Broome) in correctly identified specimens of M. brevidactylus for the north-west of Western Australia across a wide comparison. As previously mentioned, Stimpson's latitudinal range of 9°, a distance of circa 1000 km. holotype of M. brevidactylus was destroyed by fire in Specimens examined were morphologically consistent 1871, however, the Australian Museum provided throughout the study area signifying that only one specimens of M. brevidactylus from Subig Bay, Luzon in species waj present. Five adult male specimens from the northern Philippine Islands. These were identified by three sites (Monkey Mia, King Bay and Broome) selected Rathbun using Stimpson's identification criteria and to cover the coastal range, were then compared with five included the adult male specimen which McNeill (1926) M. longicarpus and three M. brevidactylus adult male described as M. longicarpus var. brevidactylus Stimpson, specimens obtained from the Western Australian in his revision of the Family Mictyridae. Museum (WAM) and the Australian Museum (AM), Thirty characters (Table 2) which were considered respectively. The WAM holding of M. longicarpus was useful diagnostically were selected for the biometric collected from Townsville in Queensland on the east study as a means of species determination, and these coast of Australia and accorded with Latreille's type were measured to the nearest 0.01 mm with a digital description. This last point is particularly important vernier caliper (DSE Model Q1382). Figures 2B, 2C, 2D & because WAM specimens examined in this study, which 2E show the location of anatomical features used in were collected from Western Australian localities and biometrics in this paper. Measurements of features (e.g., attributed to M. longicarpus, did not correspond to leg, eye, etc.) were obtained on the right side of the crab Latreille's description of the holotype of M longicarpus. Table 2 Diagnostic characters and other abbreviations used in this paper Character Description Abbreviation Abdominal segment 1 straight line length of l'1 abdominal segment measured along the mid-line AS1L Abdominal segment 6 length of 6lh abdominal segment measured along the mid-line AS6L Abdominal segment 7 length of 7“' abdominal segment measured along the mid-line AS7L Antero-lateral spines interspace distance between inner bases of antero-lateral spines ALSI Body Height Vertical distance between top of carapace and lowest point on abdomen BH Carapace anterior to posterior length between front and edge of posterior border of carapace CL (not including setae) Carapace lateral dimensions distance between widest point of branchial regions CW Cheliped carpus anterior to posterior straight line distance between top of wrist to distal end, along outer edge CHCL Cheliped carpus distal edge length of distal edge of wrist CHCW Cheliped dactyl length of moveable finger CHDL Cheliped palm lower margin length of palm on lower margin CHPLlm Cheliped palm upper margin length of palm on upper margin CHPLum Cheliped pollex length of immoveable finger CHPOL Cheliped palm depth of palm CHPW Cornea distance from top of eye to start of peduncle COL Cornea plus peduncle distance from tip of cornea to base of peduncle COL+PED Eye interspace Distance between outer extremities of eyes El Front lateral dimensions horizontal distance across top of frontal lobe FW Front vertical dimensions distance from the median point of the front, level with the eyes, to the FL point of the median lobe Gastric region lateral dimensions width of gastric region GW Median carapace dimensions width of middle part of carapace (mid-branchial region) MCW Posterior border setae length of setae lining posterior border PBH Posterior border lateral margin distance in a straight line from distal corner of carapace border to beginning PBL of curve around ischium of 4lh walking leg Posterior border caudal margin width of rear carapace border from comer to corner PBW 3rd maxilliped total length of 3rd maxilliped from top of merus to bottom of ischium 3MTL 3rd maxilliped merus central length of merus 3MML 3rd maxilliped ischium naked surface central length of bare ischium surface 3MILns 3rd walking leg merus length of part measured centrally 3WLML 4th walking leg dactyl length of part measured centrally 4WLDL 4th walking leg propodus length of part measured centrally 4WLPL 4lh walking leg merus length of part measured centrally 4WLML 35 Journal of the Royal Society of Western Australia, 91(1), March 2008 A granulated antero-lateral spine granular carapace granulated ridge from antero-lateral spine to branchial region \ peduncle and cornea / antennule / frpnt teral margin granulated ridge to subhepatic region front median _granulated subhepatic ridge suborbital ridge epistome straight line measurement of the longest length of carpus widest part of cheliped carpus CHCW 3MTL curved tip, curved tip, components of covered In setae covered In setoe pointed chitinous 3rd maxilliped structure covered pointed chitinous in setae — structure covered In setae rm e ru s' CHPLum spatulold concavity CHPUm (distal) linear callosity Ischium . keel spatulold concavity CHPOL basal flange tooth furrow S6 2' Ske'e^1 anat°my ofJMjc^ris and a key those features used in biometrics in this paper. A. Key features evident in the enor view, those annotated in bold print are diagnostic for Mictyris occidentalis. B. Dorsal view showing location of anatomical papTSDUClose-upTfechelipedhto Eate'|0bI‘?u 36 Unno: New species of soldier crab Table 3 antero-lateral spines and associated ridges, and a Ratios of morphological characters used in this paper convexly rounded posterior border, viewed dorsally, projecting beyond the abdomen. The chelipeds are Ratios following Ratios devised strongly granulated with well-defined granular ridges on McNeill (1926) for this paper the palm and fingers, and the dactyls of the fourth walking legs are distinctly curved. ALShPBW CHCL:CHCW CHPW:CHPLlm CHCL:ALSI Description of male holotype CHDLiCHCL CHPOL:CHPLlm CHPOL:CHCL CL:BH Body. Sub-globular (Fig. 3A), width between widest CHPW:CHPLlm 4WLML:PBL part of branchial regions less than length from front to CHPWiCHPOL 4WLML:4WLDL posterior border of carapace (CL:CW= 1.21:1); body CL:CW extends to slightly forward of bases of antero-lateral COL:COL + PED spines. Branchial regions inflated and projecting over ELPBW lower edges of carapace at bases of ambulatory limbs. ELMCW Posterior border of carapace (Fig. 3C) truncated and FW:FL produced beyond curve of abdomen, slightly convexly 3MML:3MlLns 3MTL:ALSI curved outward along its width which is slightly less 3MTL:CHPLum + CHDL than the interspace between the antero-lateral spines (ASLI:PBW=1.22:1); lateral margins are rounded and length of sides are one fifth the width of border; a fringe body, unless the feature was missing, in which case, the of long setae lines the border margin and sides with the left side was utilised. Averages were taken of the length of setae (PBH) being approximately three-quarters individual measurements for each group of crabs and of the length of the sides (PBL). these data were used to determine the character ratios Carapace: Visibly granular and not smooth to touch employed in the comparisons. (Figs. 2B & 3A). Gastric and branchial regional grooves The terminology used here follows McNeill (1926), well-defined. Branchial regions visibly granular with particularly in the terms for the cheliped where the wrist closely-set granules covering branchial region and = carpus, palm = manus, immoveable finger = pollex and forming rugae on the posterior halves. Median areas with moveable finger = dactyl. Abbreviations employed in this groups of macroscopic granules particularly on the paper are shown in Table 2. gastric region near the antero-lateral spines and on the cardiac mound which also has several rounded Use of ratios of particular morphological features tubercules; two chitinised stigmata on central posterior largely follow McNeill (1926), particularly for the ratios border of gastric region; two short, longitudinal grooves of CW:CL, all the 3rd maxilliped ratios, ALShPBW, adjacent to each branchial region with a pair of small CHPW:CHPLIm, and FW:FL. However, other ratios have tubercules in the centre of the median area; two small been devised for this paper to highlight diagnostic pits at base of cardiac mounds each adjacent to bases of characteristics of M. occidentalis (Table 3). 4,h walking legs. Sub-hepatic regions visible from above; strong granular ridge running along front of sub-hepatic regions downwards to oblate Milne Edwards openings Taxonomy above the cheliped basis on either side; macroscopic strong granular ridges extend inwards from sub orbital MICTYRIDAE Dana, 1851 border, diminishing to microscopic granules below Mictyris Latrielle, 1806 antero-lateral spines; antero-lateral spines (Fig. 2A), prominent, obtusely pointed, recurved, granular at bases Type species: Mictyris longicarpus Latreille, 1806 and on anterior faces; directed obliquely upwards and outwards; a macroscopic, strongly granular ridge Mictyris occidentalis, sp. nov. (Figures 2, 3) extends posteriorly from base of each antero-lateral spine to branchial region and another short, sparsely granular Mictyris longicarpus (pro parte). - McNeill 1926: page ridge extends downwards towards sub-hepatic region 109 ("north west coast" material) from base of antero-lateral spine; interspace between antero-lateral spines is almost equal the distance between Material examined the outside of the eyes, 1.22 times the width of the Holotype: male (CL 12.46 mm, CW 10.32 mm) WAM posterior border and almost equal the width of the space C39420, King Bay, Dampier Archipelago, Western between the mid branchial regions. Front is vertical (Figs Australia, 20° 38’ 16" S, 116° 45' 50" E (WGS84), coll. J. 2A and 3B), broad and channeled; length is four fifths of Unno, 24 Apr 2003. the width (Fig. 2E); lateral lobes broadly conversely Allotype. female (CL 10.06 mm, CW 8.15 mm) WAM curved, median lobe obtusely pointed and sides C39421, King Bay, Dampier Archipelago, Western obversely curved; margin of front is carinated and Australia, Lat. 20° 38' 16" S Long. 116° 45' 50" E (WGS84), granular; clusters of macroscopic granules occur on the coll. ]. Unno, 24 Apr 2003 epigastric ridges and the center of the median lobe. Eyes: Smaller than in the type species Diagnosis (M longicarpus); the space between the outside of the M. occidentalis is distinguished from other Mictyris eyes (Fig. 2B) slightly greater than the width of the species by a visibly granular carapace, especially on the posterior border (ELPBW = 1.14:1); cornea tipped with 37 Journal of the Royal Society of Western Australia, 91(1), March 2008 C cheliped front view 4th walkinvaiKing leg dactyl 5 mm posterior border 5 mm in profile \ posterior border first abdominal segment ventral view 5 mm 2 mm ; I f'ffTn3' AHnat°Tiual features °{the holotype and paratypes of Mictyris occidentals. A. Dorsal view of the holotype (KB5) showing Da^atvDe'?KBV|tstin1e'CaraPaCe,lt^eJ:,OS^er''0r boTdeTcmd its frin8e of setae' and the curved dactyl of the 4“' walking leg. B. Front of the FatemUoineC ^ ?Ff* faraPace' the carinated, granular margin of the front, and the granulated antero- Mrinar!?? 7 D key ffatup of U °cc'dental,s using paratypes. The illustrations show the nature of the granulated and cannated front margin; the granulated antero-lateral spines, and the granulated ridge from the antero-lateral spines to the branchial ofirolod"611 iff1? ^ 4 rlkin8 l6fi; ,he 8e°metry °f the gonopod; the features of the chelfpSSiSge on the eTtedo on the inside o "e dactvhE^l" edgefh°n *e^ide <»,immovable finger of the propodus, and a sfngle domed tooth ^ H the dactyl, the shape and setae on the I- abdominal segment (viewed ventrally); the shape and setae of the posterior border, and the posterior border viewed in lateral profile relative to the posterior part of the carapace. posterior 38 Unno: New species of soldier crab several long setae (Fig. 2A); cornea length is one third of granular on the palm but becomes smooth down the total length of cornea plus peduncle (COL:COL + PED = length of the pollex; the lower border of the palm has a 0.33:1). setae-lined microscopically granular ridge and the area between these two ridges is abundantly microscopically Epistome: With small gap between dorsal point and granulated; depth (width) of palm is almost equal to front (Fig. 2A); short transverse ridge just below dorsal length of lower margin (CHPW:CHPLlm = 0.90:1). point and area above that is microscopically granulated; Immoveable finger surface is largely smooth and is median area smooth and ventral projection is evenly defined on the lower border by a setae-lined groove and microscopically granulated; sides and point of ventral on the inner cutting edge by 9 irregular to rounded small projection flat with faintly finely serrated margins; ratio teeth proximal to the palm. Immoveable finger length is of width to length is 1: 2.63. slightly less than one and a half times the length of the Third maxillipeds: Large, their greatest length is lower border of the palm. Moveable finger has a broad, almost two thirds longer than the antero-lateral spine flat tooth on the cutting edge, appearing asymmetrically interspace (3MTL:ALSI = 1.56:1) and slightly longer than domed (longest side is towards palm), when viewed the length of the upper margin of the palm plus the from the side and possessing several faint serrations at moveable finger (3MTL:CHPLum + CHDL = 1.18:1); the the base proximal to palm; a groove along the dorsal length of the naked surface of the ischium is equal to the surface of the moveable finger is defined by two ridges antero-lateral spine interspace (Fig. 2C); the length of the running three quarters of the length of the finger, merus is slightly more than half that of the naked surface serrated to level of tooth and scarcely so thereafter; of the ischium (3MML:3MILns = 0.56:1); the upper another smooth ridge runs from base of finger and forms portion of the merus appears smooth while the lower a keel on the outer surface; moveable finger not quite as portion is granulated; the surface of the ischium is evenly long as the wrist (CHCL: CHDL = 1.26:1) but longer than granulated. immoveable finger; the ratios of wrist and moveable Abdomen: First segment has a projecting flange at the finger with respect to the immoveable finger (CHCL: articulation with the carapace, comprising slightly less CHPOL) being 1.59:1.18: 1, respectively; moveable finger than half the total length of the first segment; first more curved than immoveable finger as in typical form; segment narrower at junction with second segment and moveable finger tip meets immoveable finger slightly successive segments increase in width to the fifth below the latter's tip. segment then decrease in width; the sixth segment is Ambulatory limbs: Long and slender; merus of 4th equal in length to the rounded seventh segmeiat. pair covered with microscopic granules denser near base; Cheliped: Slender with large spine on each ischium carpus microscopically granular and propodus smoother directed forwards and inwards (Fig. 2C). Lower outer except for faint transverse furrows; dactyl short, broadly margin of each merus with one small spine on distal half triangular in cross-section, setose, distinctly curved as long as the antero-lateral spine but not as stout, with outwards on distal third and slightly flattened towards two large spinules/granules below this; outer surface of tip; ratio of merus length to dactyl length of 4th walking merus heavily microscopically granulated above and leg is 1.54:1; dactyl slightly exceeds propodus of 4th below the median area which is sparsely granulated; walking leg in length (4WLDL:4WLPL=1.26:1). Merus of inner surface of merus granulated with proximal end 3 rd pair almost as long as 1st to 5th abdominal segments. more heavily granulated; whole of merus appears faintly Meri of ambulatory limbs have finely serrated ridges on transversely furrowed. Wrist with tuft of setae on the antero-dorsal and ventral edges as well as postero- inner surface (Fig. 2D), two fifths of the way down from ventral edge; these ridges are particularly prominent and the trigonal joint with the merus; wrist frontal edge has macroscopically granular on the meri of the 3rd walking setae-filled groove defined by two granular ridges legs; the ventral regions of the meri are more densely running from level of tuft to end of wrist; broad rugose granulated than in the M. longicarpus; central areas of ridge with microscopic granules forming short transverse the meri are weakly transversely furrowed as in cheliped rows along its length defines deep longitudinal groove merus. Carpi are microscopically granular and the along front of wrist; remaining dorsal surface of wrist propodi are smoother except for faint transverse furrows. evenly covered with microscopic granules and with very Dactyls of 1st to 3rd walking legs are smooth with a central faint transverse furrows as in merus; length of wrist is ridge on outer surface. almost as long as length of 1st to 5"' abdominal segments. Gonopod: Comprises a flattened three-sided (cross- Palm with upper margin grooved; grooves filled with sectionally triangular) distally tapering shaft, with its setae, as in wrist, but edge of ridges more serrated; a ventral surface both convexty curved and parallel to the strong, scarcely granular ridge runs parallel to the front crab's sternum (Fig. 3C). Tlie edges of the triangular shaft edge of the palm; another scarcely granular ridge runs are keeled, and each face of the triangular form may have obtusely from the moveable finger across to the lower a longitudinal, length-parallel, broad and shallow runnel, palm edge near the junction with the wrist where it the most prominent being on the dorsal-lateral face. The almost joins with a more strongly granular narrow ridge basal end of the gonopod has a mesial tapering flange. that runs longitudinally down the length of the palm and The shaft tapers in the distal half and curves at the tip the immoveable finger; the concave area of the palm (termed here the distal curvature). The tip is torsioned delineated by these ridges is rugose with transverse (or twisted) at almost 45° out of the plane of the dorsal microscopically granular ridges and has the appearance orientation resulting in the point of the gonopod facing of faint transverse furrows; another ridge with a line of laterally outwards and parallel to the sternum. The distal short setae, near the lower edge of the palm, runs the curvature is a slightly overturned hook with a linear length of the palm to become part of the lower border of callosity on the ventral side adjacent to the sternum. A the immoveable finger; this ridge is microscopically tuft of long setae arises from the rounded end of this 39 Journal of the Royal Society of Western Australia, 91(1), March 2008 latter structure. The lateral surface of the shaft below the with convexly curved lateral margins and obtusely distal curvature is spatuloid, i.e., there is a spoon-like pointed median lobe, margins are carinated and concavity on the shaft leading up to the inner curvature. granular; 3rd maxillipeds granulated with merus slightly The extreme distal tip of the gonopod has a pointed greater than half the length of the ischium naked surface; triangular chitinous structure. The distal extremity of the abdomen with first segment flanged at posterior gonopod also is variably fringed by setae, dense enough articulation with carapace, 6"' segment is equal in length to completely cover and conceal the chitinous structure. to 7lh segment; cheliped slender, visibly granulated, with Lines of setae also may be sparsely distributed on the well-defined grooves and granulated ridges; single angular edges of the shaft, or within the length-parallel forward-facing spine on cheliped ischium; single spine runnel. Gonopod length extends from 3rd to 6th segment on merus outer margin; length of wrist is almost as long with tip being visible just above end of sixth segment. as length of 1st to 5lh abdominal segments; palm with Colour. When alive, body is coloured sky blue well-defined granular ridges; depth (width) of palm is dorsally, on the 3rd maxillipeds and on the abdominal almost equal to length of lower margin; immoveable segments; branchial regions paler with pinkish tinge; all finger length is slightly less than one and a half times the limbs, including chelipeds are uniformly cream coloured, length of the lower border of the palm; moveable finger with no colour patches on the joints. Preserved specimens not quite as long as the wrist; prominent asymmetrical still retain some dark blue colouration on the dorsal broad, domed, largely smooth tooth on cutting edge of surface and abdominal segments but otherwise are white moveable finger; merus of 3rd pair of walking legs almost except for small dark tips on the fingers. There is some as long as l'1 to 5"’ abdominal segments; dactyl of 4"' pair colour variation with adult males from Broome having of walking legs short and distinctly curved along distal orange coloured limbs and setae and specimens from half. Monkey Mia, Shark Bay having salmon pink branchial regions. All crabs examined in the study area exhibited Description of female allotype the characteristic of uniform colouration of limbs, and Material examined: Allotype: female (CL 10.06 mm, absence of colour patches on the limb joints. CW 8.15 mm) (VCSRG KB12 swarm) King Bay, Dampier Archipelago, Western Australia, 20° 38’ 16” S, 116° 45’ 50" Size: The largest adult male measured in this study E (WGS84), coll. J. Unno, 24 Apr. 2003. was from Monkey Mia, Shark Bay, with a CL of 17.5 mm from the WAM C8038 collection dated 25lh August 1960. Carapace: Sub-globular and granulated, with The smallest adult male or sub-adolescent (as defined by projecting, rounded posterior border as in male; CL:CW blue colouration and participation in swarming) from ratio 1.22:1; anterior spines smaller but more heavily SetUers Beach near Cossack, had a CL of 7.5 mm, granulated than male holotype with a row of short representing a size range of 10.0 mm. bristles running along ridge to branchial regions and a tuft of long setae on short ridge to sub-hepatic region, Variation: Large, older adult males have a more fhird maxillipeds as in male except ischium appears sculptured carapace, distinctly mounded cardiac hump more granular and evenly covered with short bristles. and robust chelipeds and ambulatory limbs. The shape of the tooth on the moveable finger may vary with age Cheliped: More slender than in male; carpus with from very asymmetric and incisor-like to a more longer setae on anterior margin; palm with ridges as in rounded, domed profile. The number of teeth on the male; fingers slender and lacking tooth on moveable cutting edge of the immoveable finger is variable, with finger and ridges on immoveable finger. 7-9 observed in the paratypes. There is some slight Walking legs. More slender than in male; dactyl on 4lh variation in the shape of the front with lateral margins walking leg curved as in male. more or less convex and centre of median lobe more or less obtusely pointed. In juvenile males, the carapace is Abdomen: Similar to male but more rounded in profile more globular, regional grooves less distinct, chelipeds resulting in a greater body height (BH); male CL:BH ratio slender with spine on merus poorly developed or absent, = 1.18:1; female CL:BH ratio = 1.11:1. The seventh and tooth on moveable finger smaller, or absent; colour abdominal segment is circa 10 % wider than in the male. brown, becoming bluer at sub-adolescence. Sexual Gonopores. Located mesially on third sternal segment; dimorphism occurs with females - they are smaller than small, simple dentoid spur projecting from end of third males of same age, have no tooth on cheliped moveable abdominal segment. finger, have obsolete spine on ischium, and their spine on the cheliped merus is reduced. Colour. Allotype specimen is bluish-brown. Adult females in life show more colour variation than adult Summary of morphological description males with some females bright blue as in the males but many are brown or grey depending on stage of moulting. Carapace visibly granular; rounded posterior border (viewed dorsally) projecting beyond abdomen; branchial Size range: Adult females are always smaller than regions uniformly granular, slightly inflated and males of the same age, being approximately three- overhanging base of ambulatory limbs; average carapace quarters the size of their male counterparts. length 13 mm, CL:CW 1.21:1; antero-lateral spines Largest female: CW = 12.5 from Shark Bay near prominent, recurved, broad, granular; prominent Carnarvon Racecourse. granular ridges from base of antero-lateral spine one extending posteriorly to branchial regions, and another Smallest female: CW = 7.1 from Settler's Beach, Cape Lambert. extending downwards to sub-hepatic regions; eyes medium, cornea not greatly wider than peduncle and one Habitat third total eye and peduncle length; front granulated Low to medium energy wave-agitated environments 40 Unno: New species of soldier crab with fine to medium (to coarse), sandy to slightly muddy taxonomically confused in the past, i.e., M. longicarpus sand substrates. Gently sloping intertidal shore zones and M. brevidactylus. M. livingstonei and M. platycheles including low gradient beaches, high to mid tidal sand have limited distributions on the east coast of Australia flats, mid tidal shoals, tidal creek banks and tidal creek and, in addition, are quite dissimilar to M. occidentalis, shoals. Usually in conjunction with a source of organic M. longicarpus and M. brevidactylus in numerous material such as mangroves or seagrass banks. morphological characters that have been well described •I in the literature (Milne Edwards 1852, McNeill 1926, Behaviour Takeda 1978). M. occidentalis does exhibit some A benthic, cryptic species, generally inhabiting the characters which appear similar to those ascribed to substrate to a depth of 10-15 cm although crabs will either M. longicarpus or M. brevidactylus, however, rapidly move deeper if excavated. Females are more closer examination shows distinct differences. For cryptic than males, rarely appearing on the surface. example, both M. longicarpus and M occidentalis have Subterranean feeding and emergence or re-entry projecting carapace posterior borders when viewed activities by crabs at low tide produces mounds, tunnels, dorsally, but are morphologically very different when and rosettes of worked substrate on the tidal-flat surface, viewed from a ventral or lateral aspect; also, and exit holes. Large numbers of crabs (predominately M. occidentalis and M. brevidactylus have curved 4th adult males) may emerge at low tide to commence walking leg dactyls, however, in the former, almost the surface feeding and thus, although crabs are behaving whole length of the dactyl is curved while in the latter, individually, may have the appearance of an "army". the curve is mainly at the tip of the dactyl. Biometric data Crabs turn sideways and burrow in a circular motion were useful in separating the new species from into the sand using the cheliped and ambulatory limbs, congeners, supporting the descriptive analysis. never the 3rt1 maxiliipeds. Some crabs, if disturbed on the The three Mictyris species of this study are surface during feeding, will run with sand packed inside systematically compared in all major morphological and extending the 3rd maxiliipeds. features in the text below and in Tables 4-6 and Figures 4-7. A summary of major morphological differences is Distribution presented in Table 7. The rationale for ordering the Sub-tropical to tropical. Recorded on the mainland species along the x-axis in Figure 7 is based on the coast of Western Australia from Monkey Mia, Shark Bay biogeographical arrangement of the data from the 25° 47' 36" S, 113° 43' 18" E (WAM C8038) to One Arm various Western Australian sampling sites which are Point, Dampier Peninsula 16° 26' 18" S, 123° 03' 42" E ordered left to right latitudinally from Shark Bay in the (WAM C39414). Also recorded within this latitudinal south to the Broome region in the north (subtropical to range from offshore islands such as Montebello Is tropical), effectively spaced in relation to their relative (McNeil, 1926) and Legendre Is, Dampier Archipelago distance from each other. This logically would place data (WAM C39408). I have examined specimens of Mictyris on M. brevidactylus, which geographically occurs in the from the Western Australian Museum holdings, collected Southeast Asian region, at least 1000 km further north of further north than One Armed Point in Western the Broome region, even further to the right on the x- Australia and from the Northern Territory region (north¬ axis, and the same distance from Broome that Shark Bay west of Unwins Island, North Kimberley, WAM C19U6, is from Broome to the south. In this framework, since Lee Point, Darwin, WAM C39389, and West Woody M. longicarpus occurs along the eastern Australian coast Creek, Gove, NT, WAM C13033) and these specimens at latitudes similar to Shark Bay in the south to Broome appear to be one species, diagnostically with two large in the north, the occurrence of M. longicarpus on the axis spines on the inner merus of the cheliped, which does of Figure 7 logically should be placed over the same not accord with the characteristics of M. occidcntalis or interval as M. occidentalis. However, since it does not M. longicarpus. They may be the "Northern Territory" form part of a geographic (climatic) gradient, and occurs species referred to by McNeill (1926), and noted by Davie in a separate geographic location, M. longicarpus is (2002) as being undescribed. No reliable records exist of placed on the far left of the x-axis. It is also placed to the soldier crab occurrence south of Shark Bay. McNeill far left to test the idea that M. occidentalis is intermediate (1926) cites a record of Mictyris from the Swan River in between M. longicarpus and M. brevidactylus. Perth by Miers in 1884 but this is highly unlikely due to the lack of suitable climate and habitat. Detailed comparison of diagnostic characters Body: There is a striking difference in body size Etymology between M. longicarpus and the other two species, the The species name is derived from the Latin former being approximately 1.5 times the latter. occidentalis meaning "of the West", alluding to the M. longicarpus is generally 25-30 mm in carapace length, location of the species in Western Australia and while M. occidentalis and M. brevidactylus are generally distinguishing it from eastern Australian congeners. only 12-15 mm in carapace length. The largest M. occidentalis recorded in this study, from examining Morphological comparison of M. occidentalis with M. thousands of individuals along the Western Australian longicarpus and M. brevidactylus coast, were ten individuals of size circa 17 mm from Monkey Mia. In describing M. occidentalis, emphasis has been placed on clearly distinguishing the new species from McNeill (1926) attributes the relatively small size of other described Mictyris species that may appear M. longicarpus var. brevidactylus (what is now superficially similar, which may have adjoining generally considered to be M. brevidactylus) to general distribution boundaries, and which have been reduction in size of species in the tropical regions. 41 Journal of the Royal Society of Western Australia, 91(1), March 2008 However, this does not explain the consistently greater than the other two species, especially on the epigastric size of the AT. longicarpus specimens from Townsville, ridges and the front margins (Fig. 4A). Queensland, a location which is definitively in a tropical All three species are divergent in region. Third maxillipeds: the relative proportions of the 3rd maxilliped merus and All three species examined in this study are sub- ischium, i.e., the length of the merus in relation to the globular in body shape, with inflated branchial regions length of the naked surface of the ischium. The merus is and truncated posterior borders. However, there is a slightly greater than half the length of the naked surface difference between the three species in the degree of of the ischium in AT. occidentalis, equal in inflation of the branchial region in relation to the base of AT. brevidactylus and slightly less than half in the walking legs: that of M. longicarpus does not overlap AT. longicarpus. The 3rd maxilliped is also the base of the walking legs, that of M. occidentalis macroscopically granulated in AT. occidentalis and slightly overlaps, and that of AT, brevidactylus distinctly microscopically granulated in AT. longicarpus and overlaps, and thus the latter has the most globular shape. AT. brevidactylus. This trend is shown in Figure 7A where M. occidentalis Abdomen: The size and morphology of the first is closer to M. longicarpus than AT. brevidactylus in the abdominal segments of all three species are different CL:CW ratio. (Figs. 5D-5F). Since they underlie the posterior border, M. brevidactylus appears to have the broadest gastric the differences of the first abdominal segments reflect the width (GW) of the three species, followed by markedly differing posterior carapace borders of the M. occidentalis and then AT. longicarpus which has the three species. All the species have an upwards slope in narrowest gastric region. This observation is the posterior part of this first abdominal segment. substantiated by the antero-lateral spine interspace to However, in the posterior part towards the carapace posterior border width ratios (Fig. 7B). The posterior border, AT. occidentalis has a broad (almost half the border of the carapace, as viewed dorsally, is length of the first segment) flat flange or platform significantly different in all species. AT. occidentalis has projecting more horizontally outwards than downwards the most noticeable posterior border which is a distinctly (Fig. 5D). The carapace also projects outwards more than horizontally projecting and convexly rounded structure, downwards to meet the abdominal segment with a minor densely fringed with long setae; AT. longicarpus has a less vertical overhang (Fig. 3C). In AT. longicarpus, on the projecting, downward-oriented, nearly straight border, other hand, the first abdominal segment slopes linearly densely fringed with short setae and M. brevidactylus upwards for the entire length of the segment, without a has no projection and a straight-edged border with a very distinct platform, to join the carapace; the lower portion short fringe of setae (Figs. 4B, 6A and 7F). of the carapace forms a vertically-long overhang, Carapace and antero-lateral spines and ridges: projecting slightly out and more vertically downwards to M. occidentalis is distinctive in its carapace, having a connect with the first abdominal segment (Fig. 5E). visibly more granular carapace that is not smooth to the AT. brevidactylus has a first abdominal segment with a touch; the other two species have a less granular short, strongly curved upward slope that meets the carapace. AT. brevidactylus is more granular in the carapace which has a minuscule overhang (Fig. 5F). median area of the carapace than AT. longicarpus. The Chelipeds: AT. occidentalis differs in cheliped branchial regions are differentially granulated in all the morphology from the other two Mictyris species in five species, with AT. occidentalis having evenly spaced, aspects: 1. a greater degree of granulation on the outer single macroscopic granules, AT. brevidactylus having surface of the wrist and the inner margins of the merus; macroscopic granular rugae and AT. longicarpus having 2. the length of the wrist is shorter than AT. longicarpus microscopic closely set granules. The antero-lateral spines (AT. longicarpus is well-named), but longer than and associated ridges (posteriorly to branchial regions AT. brevidactylus; 3. the width of the wrist is more and downwards to sub-hepatic regions) are diagnostic in slender than both species (Fig. 7E); 4. the palm is less M. occidentalis as they are visibly granular and wide, and 5. the length of the moveable finger is shorter. prominent (Fig. 5A). Of the three species, the structure of the cheliped of AT. occidentalis most resembles that of AT. longicarpus in In AT. longicarpus, the antero-lateral spines are prominent but have few granules at their bases and the the proportions of the immoveable finger, (Table 4) and ridges are ill-defined and only microscopically granular the shape of the tooth on the moveable finger. While while AT. brevidactylus has less prominent and granular AT. longicarpus is similar to AT. brevidactylus in three of anterodateral spines and the associated ridges are their cheliped ratios (1. the cheliped width is twice the significantly less granulose (Figs. 5B and 5C). wrist length; 2. the moveable finger is almost equal to the length of the wrist; and 3. the width of the palm is equal Front and eyes: AT. longicarpus is distinctive in ha vim to the length of the lower palm border), the cheliped large globulose eyes on stout peduncles (Fig. 5B) and a ratios of AT. occidentalis are dissimilar in all respects to narrow-appearing front that has nearly straight lateral those of AT. brevidactylus. Significant ratios of cheliped margins with the front width being equal to front length, features are presented in Table 4 and Figure 7E and In contrast, AT. occidentalis and AT. brevidactylus have chelipeds of the three species are illustrated in Figure 4D. smaller eyes, but similar in size to each other. However, In adult males, the tooth on the moveable finger is M occidentalis has a narrower front (front length is four asymmetrically domed in AT. occidentalis, conical in AT. ‘ ^S . ,fr°n! w'dth) than M. brevidactylus. longicarpus and asymmetrically serrated in AT M. brevidactylus has the broadest front (front length is brevidactylus (Fig. 4D). wo thirds front width) of the three species (Fig. 7D). Walking legs: All three species show similarity in the M occidentalis exhibits greater granularity in its front slenderness of the walking legs. The most useful 42 Unno: New species of soldier crab Figure 4. Comparative anatomy of Mictyris occidentalis, M. longicarpus, and M. brevidactylus. in each diagram, for M. occidentalis, individuals from Shark Bay, King Bay (Dampier Archipelago), and Broome are illustrated to show intra-species and geographic variation. A. The anterior front view of the three species showing: shape of front and the extent of granulation along its edge; eyes and relative size of cornea and peduncle; antero-lateral spine and the extent of its granulation; and the extent of granulation of the dorsal carapace; inset illustrates the differences between the three species by focusing on the shape of the front, and the orientation and extent of granulation of the antero-lateral spine. B. Shape of the posterior border, and extent that the posterior border extends past the point of insertion of the 4"’ walking leg. C. Dactyl of the 4lh walking leg: M. occidentalis is relatively short and curved, M. longicarpus is relatively long, and M. brevidactylus is short and curved but more slender than that of M. occidentalis. D. Chelipeds: M. occidentalis has a serrated edge on the inside of the immovable finger of the propodus, and a single domed tooth on the inside of the dactyl; M. longicarpus also has a serrated edge on the inner surface of the immovable finger of the propodus, and a single conical tooth on the inside of the dactyl; M. brevidactylus lacks a serrated edge on the inside of the immovable finger of the propodus, and has an asymmetrical blade-like, triangular single tooth on the inside of the dactyl that is serrated on its proximal edge and smooth on its distal edge; inset diagrammatically illustrates the differences in the tooth on the inside of the dactyl for the three species. 43 Journal of the Royal Society of Western Australia, 91(1), March 2008 2 mm 4 M. brevidactylus Io(nt‘lrpus' and M. brevidactylus. A-C. Dorsal view of carapace front for peduncle, tl/size of the eves ind the vS™ ?ranulat,°" of the carapace granulation of the front, the length and thickness of the showing thevariat^on^in'itTplan^shape between'the'^speciM^he^extent'thatch’Dr^s8^'1165 WeW \**"*•' «*“«* of development of setae along the posterior margin. F H dLPressions and ndSes comprise the form, and the extent diagnostic features of the legs for comparative purpo: greater curvature (Fig. 4C and Fig. 7C). M. occidentalis are the dactyl of the 4lh walking leg with respect to and M. brevidactylus are similar in having the length of shape and length in relation to the propodus of the sat the 3rd WL merus equal the length of the lsl to 5th limb, and the merus of the 3rd walking leg with respect abdominal segments, while that of M. longicarpus equals its total length. M. longicarpus is easily distinguish the lsl to 6th abdominal segments. from the other species, with its long, relatively straw Gonopods: The gonopods of Mictyris examined in this 4"’ WL dactyl (4WLDL: 4WLML = 1.22: 1). M. Occidenta study, though broadly similar, show some discernible and M. brevidactylus both have short, curved 4lh V differences between the species. A gonopod of dactyls, however, M. brevidactylus is slightly the shorl M occidentalis is illustrated in Figure 2F. The key (1.65 times the merus length as opposed to 1.50 times features that help to distinguish between the three M. occidentalis) and the dactyl of M. occidentalis has species are: beginning point of torsion on shaft (viewed 44 Unno: New species of soldier crab location of the posterior border profiles illustrated adjacent B Gonopods 1 mm chitinous structure distal linear 0 callosity keel M. occidentalis M. longicarpus M. brevidactylus Figure 6. Comparative anatomy of Mictyris occidentalis, M. longicarpus, and M. brevidactylus. A. Variation in the profile shape of the posterior border. Cross section of carapace shows where the detailed drawings of the posterior border are located. Individual drawing shows a prominent sloping platform for the posterior border for M. occidentalis, a steep, relatively high protrusion for M. longicarpus, and a short sloping platform for M. brevidactylus. B. Variation in gonopod morphology between the three species. on the anterior surface); extent of curvature of the tip of shaft. A comparison of the three species is provided in the shaft; length of the chitinous structure relative to the Table 5 and Figure 6B. diameter of the distal end of the shaft; extent of Of the three congeners, the gonopod of M. occidentalis development of the concavity of the spatuloid shape; the exhibits the greatest degree of torsion with the twist extent of development of the keel; and the development beginning at the midpoint of the shaft rather than higher of the longitudinal furrow. Setae and their extent of towards the distal end as in the other species. In general development did not appear to be species-diagnostic appearance, the gonopod of M. occidentalis has greater except that JVf. brevidactylus was sparsely setose on the similarity with M. brevidactylus than with 45 Journal of the Royal Society of Western Australia, 91(1), March 2008 Table 4 Comparison of cheliped morphology Cheliped ratios M. occidentalis M. longicarpus M. brevidactylus Explanation of ratios CHCL:CHCW: 2.49:1 2.01:1 1.99:1 wrist length is 2 xh times wrist width in M. occidentalis and - 2 times in M. longicarpus & M. brevidactylus CHPOL:CHPLlm 1.45:1 1.42:1 1.00:1 immoveable finger is ~ 1 Vi times length of palm lower margin in M. occidentalis & M. longicarpus but equal in M. brevidactylus CHDL:CHCL 0.82:1 0.90:1 0.94:1 moveable finger much shorter than wrist in M. occidentalis, slightly shorter in M. longicarpus and almost equal in M. brevidactylus CHPOL:CHCL 0.71:1 0.75:1 0.58:1 immoveable finger is - 3A length of wrist in M. occidentalis & M. longicarpus but - 2/3 in M. brevidactylus CHPWrCHPLlm 0.80:1 0.90:1 0.91:1 palm width < length of palm lower margin in M. occidentalis and almost equal in M. longicarpus & M. brevidactylus CHPW:CHPOL 0.64:1 0.66:1 0.75:1 palm width is V, length of immoveable finger in M. occidentalis. & M. longicarpus but % in M. brevidactylus CHCL:ALSI 1.30:1 1.51:1 1.15:1 wrist length is 1'/, times antero-lateral spine interspace in M. occidentalis, 1 Vi times in longicarpus and almost equal in M. brevidactylus Table 5 Comparison of gonopod morphology Gonopod feature M. occidentalis M. longicarpus M. brevidactylus torsion of shaft begins at midpoint of shaft begins just before curve of tip begins at three-quarters of length of shaft curvature of the tip curved with slightly curved with short hooked tip curved with distinctly extended tip of the shaft extended tip description of the long and thin long and thick short and thin chitinous structure location of the in line with the apex of below the apex of the in line with the apex of curvature chitinous structure curvature of the shaft curvature of the shaft of the shaft concavity of the well developed moderately developed weakly developed spatuloid shape keels prominent prominent present, but weakly developed furrows prominent prominent weak setae along shaft and well along shaft and well generally sparse along shaft, but well developed at tip developed at tip developed at tip length from 3,d abdominal segment from 3rd abdominal segment to from 3rd abdominal segment to just above to just above 6,h segment below top of 6lh segment (not 6,h abdominal segment (visible) (visible) always visible) Table 6 Comparison of gonopore morphology Gonopore feature M. occidentalis M. longicarpus M. brevidactylus size of spur small large almost obsolescent shape of spur simple, dentoid, blunt point multi furcate, dentoid, blunt point simple, sharp point setae sparse fringe dense short fringe at end very sparse 46 Unno: New species of soldier crab Figure 7. Ratio of various anatomical features of Mictyris occidentalis, M. longicarpus, and M. brevidactylus. The grey band in each graph shows the variation in value of the ratio for the holotype and the 14 paratypes of M. occidentalis, based on three latitudinally widespread sites, with 5 male crabs from each site. A. The ratio of carapace length to carapace width which shows M. longicarpus and M. brevidactylus are just outside the degree of variation of the ratio for M. occidentalis. B. The ratio of antero-lateral spine interspace to posterior border width with M. longicarpus and M. brevidactylus just outside the degree of variation of the ratio for M. occidentalis, and with M. occidentalis intermediate in value to those of the other two species. C. The ratio of 4th walking leg merus length to 4th walking leg dactyl length, with M. longicarpus and M. brevidactylus definitively outside the degree of variation of the ratio for M. occidentalis, but with M. occidentalis intermediate in value to those of the other two species. D. Tire ratio of front width to front length with M. longicarpus and M. brevidactylus definitively outside the degree of variation of the ratio for M. occidentalis, but again with M. occidentalis intermediate in value to those of the other two species. E. The ratio of cheliped carpus length to cheliped carpus width with M. longicarpus and M. brevidactylus definitively outside the degree of variation of the ratio for M. occidentalis, but with M. occidentalis not intermediate in value to those of the other two species. F. The ratio of 4th walking leg merus length to posterior border length; M. longicarpus is similar to M. occidentalis, and M. brevidactylus definitively outside the degree of variation of the ratio for M. occidentalis. 47 Journal of the Royal Society of Western Australia, 91(1), March 2008 M. longicarpus in that the head of the distal curve is Species Key longer, although not to the extent of M. brevidactylus. However, the shaft has more affinity to M. longicarpus A key to distinguish the adult males of the five with prominent keels and furrows. described species of Mictyris Latreille, 1806 including the three species considered in this paper, is presented A summary of the major morphological similarities below: and differences of the three Mictyris species considered in this study is presented in Table 7 below. Key to Mictyris Latreille, 1806 Gonopores: Gonopores of the three species share the Al. Posterior border of carapace conspicuously same feature common to the genus of being a cavity produced and overlapping abdomen overhung on the mesial side with a projection or spur Bl. Carapace macroscopically granular, antero-lateral formed from the end of the 3rd abdominal segment. The spines and ridges prominent, and strongly greatest variation in the gonopores is in size and granular, branchial regions overlapping base of complexity of the spur ranging from an almost legs, dactyl of 4th WL distinctly curved along distal obsolescent, pointed spur in M. brevidactylus, to a small, third, average size, 12 mm CL.occidentals blunt spur in M. occidentals, to a large, complex spur in M longicarpus (Table 6). Table 7 Summary of major differences between Mictyris species Species Character M. occidentalis M. longicarpus M. brevidactylus carapace macroscopically granular, not appears smooth, microscopically sparsely granulated, not entirely smooth to touch granular, smooth to touch smooth to touch branchial regions moderately inflated, slightly slightly inflated, not overlapping distinctly inflated, overlapping base of overlapping base of walking legs; base of walking legs; walking legs; macroscopically granular macroscopically granular with microscopically granular with rugae single granules posterior border distinctly projecting and convex projecting, slightly sinusoidal not projecting, straight antero-lateral broad and granular, recurved, prominent, recurved, oriented prominent, recurved, oriented upwards spines and ridges oriented outwards more than outwards more than upwards; more than outward; sparsely granular, upwards; prominent granular smooth except for sparse sparsely granular ridge running ridge running backwards to granules at base; ill-defined ridge backwards to branchial regions and branchial region and another running backwards to branchial another short, microscopically granular short, strongly granular ridge regions and another short, ridge downwards to sub-hepatic region downwards to sub-hepatic microscopically granular ridge region downwards to sub-hepatic region front broad, convexly curving laterally narrow, straight laterally very broad, straight laterally eyes medium, globular large and globulose; stout small, globular peduncle third maxillipeds lower part of merus and all of microscopically granulated; microscopically granulated; length of ischium heavily granulated; length of merus less than half merus half that of naked surface of length of merus slightly more that of naked surface of ischium ischium than half that of naked surface of ischium Abdomen 7"' segment equal to 6,h 7th segment longer than 6,h 7"’ segment equal to 6,h Chelipeds wrist and palm slightly more Wrist and outer surface of palm carpus and palm wide; fingers short slender; front surface of wrist smooth; fingers long and robust- (immoveable finger almost equal and margins of merus visibly large conical tooth on moveable length of lower palm) and sturdy; granulated; narrower palm and finger broad, serrated asymmetrical tooth on shorter moveable finger; fingers moveable finger tapering at ends; slightly asymmetrical domed tooth on moveable finger with faint serrations on side proximal to palm walking legs merus of 3rd pair as long as Is' to merus of 3rd pair as long as l5' merus of 3rd pair as long as I5' to 5th 5,h abdominal segments; dactyl of to 6,h abdominal segments; abdominal segments; dactyl of 4lh 4'" walking leg short, setose and dactyl of 4lh walking leg long walking leg short and distinctly curved distinctly curved over distal third and slightly curved at tip at tip gonopods sturdy shaft, slender tip stout, stout, short distal curve, slender, very long distal curve, short keeled, long distal curve, prominent chitinous tip chitinous tip prominent chitinous tip gonopores small, simple spur large, setose, multifurcate spur almost obsolete simple spur 48 Unno: New species of soldier crab B2. Carapace not macroscopically granular, appearing walking leg merus length to 4th walking leg dactyl length smooth; branchial regions not overlapping base of (Fig. 7C), and the front width to front length (Fig. 7D), walking legs; antero-lateral spines recurved, M. occidentalis indeed appears to be intermediate without strongly granular ridges, dactyl of 4th WL between M. longicarpus and M. brevidactylus. However, straight with slightly curved tip; large globulose for the ratios of cheliped carpus length to cheliped carpus eyes, large size, 20-30 mm CL.longicarpus width (Fig. 7E) and 4U' walking leg merus length to A2. Posterior border of carapace produced but not posterior border length (Fig. 7F), M. occidentalis does not overlapping abdomen; carapace covered with appear to be intermediate between M. longicarpus and whitish tubercules, branchial regions dorsally M brevidactylus. Similarly, other morphological features swollen, average size 16 mm CL.platycheles of the three Mictyris species do not support the idea that M. occidentalis is an intermediate between A3. Posterior border of carapace not conspicuously M. longicarpus and M. brevidactylus. These include, for produced instance, some of the morphological features of the Bl. Prominent broad serrated tooth on cutting edge of gonopods (Table 5), as well as the ratio of the cheliped moveable finger; granular ridge from antero-lateral carpus length to cheliped carpus width, the ratio of the spine posteriorly to branchial region, average size 4th walking leg merus to the posterior border length the 12 mm CL.brevidactylus shape of the posterior border, the shape of the Is1 abdominal segment, and the degree of granularity of the B2. Obsolete tooth on cutting edge of moveable finger; carapace, the morphology of the antero-lateral spines, the no ridge from antero-lateral spine posteriorly to shape of the front, and the granularity of the 3rd branchial region, small eyes, average size, 11 mm maxilliped. CL.livingstonei In summary, while M. occidentalis shows some morphological similarity to M. longicarpus, and another set of similarities to M. brevidactylus, it has some of its Discussion and conclusions own unique features. 1 consider it not to be an Qualitative and biometric data point to the Western intermediate between M. longicarpus and Australian form of Mictyris as a new species. It is M brevidactylus. morphologically distinct, and can be readily The focus on comparing the morphology of the three distinguished from M. longicarpus by its smaller size, species in this study has provided interesting information rounded and projecting posterior border, more granular as to where the major variations in the characters of the carapace, more inflated branchial regions, prominent genus occurs. These characters are: the arrangement of ridges extending from the antero-lateral spines, broader the carapace posterior border-first abdominal segment front, and short, curved dactyl on the 4th walking leg; interface; the degree of inflation of the branchial region; and from M. brevidactylus by its projecting posterior the extent of granulation on the carapace; the form and border, more granular carapace, less inflated branchial granulation of the antero-lateral spine; the prominence of regions, prominent ridges extending from the antero¬ the branchial and sub-hepatic ridges extending from the lateral spines, narrower front and domed tooth on antero-lateral spines; the shape and size of tire tooth on moveable finger of cheliped. the moveable finger; and the shape and size of the dactyl The ratios of morphological features of M. longicarpus of the 4th walking leg. They appear to be the characters and M. brevidactylus determined in this study that consistently vary, though in different forms, from corroborate the ratios for these species determined in species to species. Of these, I consider that the earlier studies by Latreille (1806), Stimpson (1858), and arrangement of the carapace posterior border - first McNeill (1926). Many of the ratios appear to be species- abdominal segment interface to be the most significant, diagnostic (viz., the ratios of carapace length to carapace and hence it is used as the first morphological criterion width, antero-lateral spine interspace to posterior border in the key to differentiate the species. width, front width to front length, and 4th walking leg On the other hand, there are numerous other merus length to dactyl length) and are significantly morphological features that do not vary markedly different between all three species. Additionally, between species. Setting aside the tooth on the moveable M. occidentalis is clearly distinguished by the ratio of finger, the shape of the chelipeds and the general pattern cheliped carpus length to cheliped carpus width, while of the cheliped ridges and grooves appear to be relatively the ratio of the merus length of the 4"' walking leg to the consistent unlike the major cheliped of the Ocypodid crab posterior border length discriminates M brevidactylus Uca which can vary greatly in form between species from the others (Fig. 7). (Crane 1975; George & Jones 1982). Similarly, the general Some authors, (e.g., McNeill 1926), have considered structure of the abdominal segments (except for the lsl the Western Australian form of Mictyris (M. occidentalis segment), and the T1 to 3rd walking legs seem to be of this paper) to be a morphological intermediate similar between the three Mictyris species of this study. between M. longicarpus and M. brevidactylus, and to test As such, these features are not afforded significant status this idea, ratio data are presented in graphical form (Fig. in differentiating between the species. 7) with data on M. occidentalis placed on the x-axis In terms of biogeography and regional species between the two congeners. Placement of the reduced variation, the graphs of ratios of morphological features data of ratios in this order in Figure 7 illustrates some in Figure 7 highlight some interesting patterns. While significant patterns: for the ratios of carapace length to there appear to be trends in ratios between the three carapace width (Fig. 7A), the antero-lateral spine congener species, as discussed earlier, there is a relative interspace to posterior border width (Fig. 7B), the 4lh consistency in results for M. occidentalis for five ratios 49 Journal of the Royal Society of Western Australia, 91(1), March 2008 across a latitudinal spread of 17°S to 25°S (a distance of De Haan 1935 ?Ocypode (IVb'ctyris) deflexifrons. Seibold's Fauna circa 1000 km), supporting observational data for a Japonica, Crustacea 2, p 25 (sine descr.). monospecific distribution from Shark Bay to Broome, and De Man J G 1887 IMytiris longicarpus. Archiv fur indicating for most of the morphological features, little Naturgeschicthe Iiii, I, p 358. variation across the biogeographic range of the species. George R W & Jones D S 1982 The Fiddler Crabs of Australia As such, any morphological deviation of M, occidentalis (Ocypodinae: Uca). Records of the Western Australian Museum Supplement No. 14. Western Australian Museum, from the other congeners becomes significant. The Perth. nearest occurrence of a described congener to Jones D S 2004 The Burrup Peninsula and Dampier Archipelago, M. occidentalis is M. brevidactylus and the Western Australia: an introduction to the history of its morphological features for M. occidentalis and discovery and study, marine habitats and their flora and M brevidactydus reflected in the ratios of Figure 7D-7F fauna. Records of the Western Australian Museum are relatively consistent for M. occidentalis over its Supplement No. 66: 27-49 Western Australian Museum, geographic range, but definitively different for Perth. M. brevidactylus over the same geographic distance, thus Kawaguchi S 2002 Effect of tube-type burrow by soldier crab corroborating that there are two separate forms. Mictyris longicarpus var. brevidactylus on alteration of soil microflora in the tidal flat of mangrove forest. Mangurobu ni Even putting aside their reproductive morphological kansuru Chosa Kenkyu Hokokusho. Heisei 13 Nendo. differences, given the fact that populations of Kemp S 1919 Notes on Crustacea Decapoda in the Indian M. brevidactylus are some 1000 km further north of any Museum. XII. Scopimerinae. Records of the Indian Museum M. occidentalis populations, and that the nearest 16: 305-348 pis. 12 -13. occurrence of M. longicarpus is several thousand Latreille P A 1806 Genera crustaceorum et Insectorum kilometres to the east in Queensland, these three secundum ordinem naturalem in familias disposita. Parisiis congeners, occurring in separate, non-overlapping et Argentorati. Vol I xviii 302 pp pis 1-16 [21] [gender geographic areas, clearly are unable to cross-breed, and masculine]. are therefore allopatric. In this context, M. occidentalis, is Latreille P A 1829 Mictyris longicarpus. In: Cuvier's Regne geographically limited to the northwest coast of Western Animal, 2nd edition 4: 47. Australia and hence is endemic. McNeill F A 1926 A Revision of the Family Mictyridae. Studies in Australian Carcinology No. 2. Records of the Australian Museum 15:100-128, pis. ix-x. Acknowledgements: Dr Diana Jones and staff of the Western Australian Miers E J 1884 Crustacea. In: Report of the zoological collections Museum were very helpful in providing specimens for examination as made in the Indo-Pacific Ocean during the voyage of HMS was the Australian Museum which provided M. brevidactylus material. "Alert", 1881-2. The author is grateful to the V & C Semeniuk Research Group for providing funding for this research, and thanks Dr Vic Semeniuk and Milne Edwards H 1852 De la famille des ocypodides Associate Professor Adrianne Kinnear for advice, support, and assistance (Ocypodidae). Second Memoire, in Observations sur les throughout this study. affinites zoologiques et la classification naturelle des Crustaces. Annales des Sciences Naturelles (Zoologie) Serie3 18: 128-166 pis 3, 4 [154], References Sakai T 1976 Crabs of Japan and the Adjacent Seas. Kodansha Ltd, Tokyo, Japan, pp 321-322, 626-627, pi 213 Fig. 1, Figs. 2, Alcock A 1900 Materials for a Carcinological Fauna of India No 3. 6. the Brachyura Catametopa or Grapsoidea. Journal of the Shih J T & Liao C F 1998 Conversion of Cholesterol to Sex Asiatic Society of Bengal 69 Pt 2 No 3: 383-384. Steroid-like Substances by Tissues of Mictyris brevidactylus Crane ] 1975 Fiddler Crabs of the World Ocypodidea: Genus in Vitro. Zoological Studies 37(2): 102-110. Uca. Princeton University Press, Princeton, New Jersey. Stimpson W 1858 Crustacea Ocypodoidea. Prodromus Dana J D 1851 Conspectus Crustaceorum qua: in Orbis descriptionis animalium evertebratorum quae in Expeditione Terrarum circumnavigatione, Carolo Wilkes e Classe ad Oceanum Pacificum Sepentrionalem, a Republica Reipublicae Foederatae Duce, lexit et descripsit. Proceedings Federata missa, Cadwaladaro Ringgold et Johanne Rodgers of the Academy of Natural Sciences of Philadelphia 5: 247- Ducibus, observatorum et descriplorum. Pars V. Proceedings 254. of the Academy of Natural Sciences of Philadelphia, 10(5):99. Davie P j F 1982 A preliminary checklist of Brachyura Stimpson W 1907 Report on the Crustacea (Brachyura and (Crustacea: Decapoda) associated with Australian Mangrove Anomura) collected by the North Pacific Exploring Forests. Operculum 5 (4): 204-207. Expedition 1853-1856. Smithsonian Miscellaneous Collections 49, No. 1717, p 103, pi 13, fig 4. Davie P J F 1985 The biogeography of littoral crabs (Crustacea: Decapoda: Brachyura) associated with tidal wetlands in Takeda M 1978 Soldier Crabs from Australia and Japan. Bulletin tropical and sub-tropical Australia Mangrove Monograph of the National Science Museum Tokyo Series A (Zoology) 4 No. 1. Darwin. In: Coasts and Tidal Wetlands of the (1): 31-38. Australian Monsoon Region (K N Bardsley, J D S Davie & C Takeda S 2005 Sexual differences in behaviour during the D Woodroffe). Australian National University North breeding season in the soldier crab (Mictyris brevidactylus). Australia Research Unit, Darwin, pp. 259-75. Journal of Zoology 266: 197-204. Davie P J F 2002 Crustacea: Malacostraca: Eucarida (Part 2): Decapoda- Anomura, Brachyura. In: Zoological Catalogue of Australia Vol 19.3B (eds A Wells & W W K Houston). CSIRO Publishing, Melbourne, xiv 641 pp. 50 Journal of the Royal Society of Western Australia, 91: 51-52, 2008 Range extension for the Black¬ headed Worm-lizard, Aprasia Notes picturata (Squamata: Pygopodidae) on .... S A Thompson1 & G G Thompson2' o 1 Coffey Environments, Dilhorn House, 2 Bulwer Street, Perth, WA, 6000 El [email protected] 2 Edith Cowan University, Joondalup Drive, Joondalup, WA, 6027 E [email protected] * corresponding author Manuscript received August 2007; accepted February 2008 Abstract. The third capture of an Aprasia picturata increases its recorded geographical distribution in a northerly direction by approximately 250 km. This record may represent a new population or a range extension. If this record indicates a small disjunct population, and the species is already relatively rare, then there may be a case for affording it and its habitat special protection. Key words: Aprasia picturata, geographic range, Western Australia Introduction The habitat of the Black-headed Worm-lizard, Aprasia picturata, was described by Wilson & Swan (2005) as Figure 1. Location of three specimens of A. picturata. 'rocky ridges vegetated with low acacias and eremophilas near Leonora, in the southern interior of rare or there have been few fauna surveys in its habitat. Western Australia' (pp 106). Given the mining 'boom' in Western Australia and the Aprasia picturata is only known from two specimens number of fauna surveys (—15) that have recently been collected at Minara Station near Leonora. Nothing is conducted in the region (ATA Environmental 2007; Biota known of its ecology (Thompson & Thompson 2006). 2006 and references therein) it is suprising that the We report here the capture of an additional specimen species has not been recorded more often, if it is approximately 30 km south of Wiluna, Western abundant. The available data would suggest that it has a restricted geographic distribution and is also rare. Australia. This represents a range extension of approximately 250km in a northerly direction The lack of records for this species makes it difficult to comment on its preferred habitat. Wilson & Swan (2005) described A. picturata habitat as being associated with Observations rocky ridges vegetated with low Acacias and Eremophilas, however, the individual reported here was On 16th January 2007, we captured an individual A. captured on a spinifex sand plain. It is presumed that A. picturata at location (WGS 84) 51 243140E, 7020828N in a picturata is similar to other Australian Apraisa, being funnel trap as part of an environmental impact fossorial and living 'beneath embedded stumps, in insect assessment for a proposed mining project in the region. holes and under rocks and logs' (Wilson & Swan 2005). The habitat comprised of a low, dense spinifex (Triodia It is not unusual to find isolated populations of reptiles sp.) grassland with occasional mixed Acacia shrubs. The outside their published geographical distribution (e.g. specimen was lodged with the Western Australian Thompson et al. 2005; Menz & Cullen 2006). However, Museum (R166877) and identified by B. Maryan. this record represents a significant extension to the published distribution (Wilson & Swan 2005; Thompson & Thompson 2006). If we adopted the precautionary Discussion principle, then there may be a prima facie case for Only two specimens of Aprasia picturata had been affording a level of special protection to this species until recorded in northern Goldfields of Western Australia. more information is available on its geographic Figure 1 indicates the recorded locations of the two distribution, population size and ecology. If there are specimens in the Western Australian museum collection only a few localities that support these disjunct and the extension to its published distribution populations of A. picturata, then appropriate habitat represented by this specimen. This may indicate that it is protection strategies should be put in place. 51 Journal of the Royal Society of Western Australia, 91(1), March 2008 Acknowledgements: The authors would like to thank Paul Doughty and Brad Maryan from the Western Australian Museum for access to the specimen database, and Cale Alexander, Carl Gosper and Edward Swinhoe for assistance with the field work. References ATA Environmental 2007 Terrestrial Vertebrate Fauna Assessment, Honeymoon Well. Unpublished Report for LionOre Australia. (Biota) Biota Environmental Sciences 2006 Fauna habitats and fauna assemblage survey of the Mt. Keith Mine Project Area. Unpublished Report for Nickel West Operations, BHP Billiton. Menz MHM & Cullen PP 2006 Occurrence of the Barking Gecko Underwoodisaurus milii (Bory 1825) (Gekkonidae) in the Pilbara Region, Western Australia. Journal of the Royal Society of Western Australia, 89: 89-90. Thompson SA & Thompson GG 2006 Reptiles of the Western Australian Goldfields. Goldfields Environmental Management Group. Kalgoorlie WA. Thompson SA, Withers PC, Thompson GG & Robinson D 2005 Range extension for the Perentie, Varanus giganteus (Squamata: Varanidae). Journal of the Royal Society of Western Australia, 88: 41-43. Wilson S & Swan G 2005 A complete guide to reptiles of Australia. Reed New Holland. 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