Aquatic fauna refuges in Marrggaret River and the Cape to Cape region of ’s Mediterranean-climatic Southwestern Province Mark G. Allen1,2, Stephen J. Beatty1 and David L. Morgan1* 1. Freshwater Fish Group & Fish Health Unit, Centre for Fish & Fisheries Research, School of Veterinary and Life Sciences, Murdoch University, South St, Murdoch 6150, Western Austrralia 2. Current address: Department of Aquattic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, , , 6986, Australia * correspondence to [email protected]

SUUMMARY and the Cape to Cape region in the extreme south-western tip of Australia are located between Capee Naturaliste in the north and Cape Leeuwin in the south and encompass all intervening catchments that drain westward to the In- dian Ocean. The region has a Mediteerranean climate and houses 13 native, obligate freshwater macrofauna species (i.e. fishes, decapod and a bivalve mol- lusc), four of which are listed as threatened under State and/or Commonwealth leg- islation. The most imperiled species are the Margaret River Burrowing ( pseudoreducta) and Hairy Marron ( tenuimanus), both of which are endemic to the Margaret River catchment and listed as critically endangered (also by the IUCN), and Balston’s Pygmy Perch (Nannatherina balstoni) which is vulner- able. The region also houses several fishes that may represent neew, endemic taxa based on preliminary molecular evidence. Freshwater ecosystems in the region face numerous threats including global climate change, a growing huuman population, introduced species, destructive land uses, riparian degradation, waater abstraction, declinning environmental flows, instream barriers, and fire. Here we review the cur- rent knowledge of the considerable aquatic values of the region to pro-

vide a contemporary checklist, and to highlight actions that may be considered to protect these values in the face of both current and future conservatiion threats.

Keywords: Threatened species; teleosts; decapod crustaceans; agnathans; mussels

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Citation: Allen MG, Beatty SJ, Morgan DL (2017) Aquatic fauna refuges in Margaret River and the Cape to Cape region of Australia’s Mediterranean-climatic Southwestern Province. FiSHMED Fishes in Mediterranean Environments 2017.002: 26p

INTRODUCTION water resources (CCG, 2005a, 2005b, 2006). In light of this, the Department of Water The south-west of Western Australia (Government of Western Australia) recently is a global hotspot of species endemism that proclaimed several unregulated water re- is influenced by a Mediterranean climate sources in the CCR for allocation under li- (Myers et al., 2000; Olson & Dinerstein, cense (DoW, 2009) and addressed the need 2002; Morgan et al., 2014a). The aquatic for water management reform in the South fauna of the region is species impoverished, West Regional Water Plan 2010 – 2030 yet the region has the highest rate of species (DoW, 2010). Balancing human water de- endemism of any major drainage division in mands with ecological water requirements Australia, with >90% of the native aquatic in a drying climatic region, where water macrofauna (i.e. fishes, decapod crusta- demand is predicted to be at the limit of ceans, and bivalve molluscs) being endemic supply within 20 years, looms as a key to this region (Morgan et al., 2011, 2014a). management challenge (DoW, 2010). More- Some of these endemic species have suffered over, the projected continuation of the cur- dramatic range declines during the past rent drying trend beyond this timeframe century, mainly due to impacts associated places further uncertainty on the long-term with deforestation and agricultural devel- of some of the region’s aquatic opment, whilst others are naturally re- ecosystems (Suppiah et al., 2007; CSIRO, stricted to narrow ranges (Morgan et al., 2009a, b; Morrongiello et al., 2011). 1998, 2011; Unmack, 2001; Allen et al., The obligate freshwater macrofauna 2002; Klunzinger et al., 2015). While land of the CCR comprises 13 native species clearing is now less prevalent, the entire (Morgan et al., 2011), four of which are regional aquatic fauna is becoming increas- listed as threatened under the Wildlife Con- ingly jeopardised by a number of anthropo- servation Act 1950 and/or Environment Pro- genic stressors and environmental threats, tection and Biodiversity Conservation not the least of which is human-induced (EPBC) Act 1999. Two additional species global climate change (Suppiah et al., 2007; are listed as ‘priority taxa’ by the Depart- Morrongiello et al., 2011; Beatty et al., 2014; ment of Parks and Wildlife, Government of IPCC, 2014; Morgan et al., 2014a). Conse- Western Australia (Morgan et al., 2011, quently, several of the region’s endemic 2014a). The most imperiled are the Hairy aquatic species are now imperiled and offi- Marron (Smith 1912) cially recognised as threatened under State and the Margaret River Burrowing Crayfish and/or Commonwealth legislation (Morgan Engaewa pseudoreducta Horwitz & Adams et al., 2011, 2014a). 2000, two range-restricted endemics that Situated within south-western Aus- are listed as critically endangered under the tralia, the Cape to Cape region (CCR) (Fig- EPBC Act 1999 (and also by the IUCN), ure 1) is globally recognised as a premier while Balston’s Pygmy Perch Nannatherina food and wine producing region, as well as a balstoni Regan 1906, and Carter’s Freshwa- popular tourist destination. The population ter Mussel Westralunio carteri (Iredale of the CCR has expanded in recent decades 1934) are either vulnerable (N. balstoni) or and land use patterns have shifted accord- qualify as vulnerable (W. carteri) under the ingly (CCG, 2005a, 2005b, 2006; DoW, 2009, EPBC Act 1999 (Morgan et al., 2014b, 2010). For instance, agriculture has intensi- Klunzinger et al., 2015). Notwithstanding fied (particularly viticulture), resulting in a the threat posed by climate change, these growing demand for surface and subsurface rare species also face a number of further 2

Threatened fauna of Australia’s Cape to Cape region 2017.002: 26 specific threats (see Burnham et al., 2012; The CCR has a Mediterranean cli- Duffy et al., 2014; Klunzinger et al., 2015). mate characterised by hot, dry summers With almost half of the CCR’s native and mild, wet winters. Long-term mean an- freshwater macrofauna recognised as nual precipitation at Margaret River is 1131 threatened or requiring conservation man- mm, and is similar at other weather sta- agement, freshwater ecosystems in the re- tions in the vicinity, but considerably higher gion have justifiably received considerable than areas to the region’s north (BoM 2015). attention from natural resource managers, Precipitation decreases to around 900 mm scientists and conservation organisations in further inland near the headwaters of the recent years. Increasing resources have Margaret River catchment (BoM, 2015). On been channelled into conservation and land- average, >75% of precipitation falls in the care initiatives in the region, such as the five months from May to September (BoM, construction of two rock ramp fishways on 2015). Evaporation rates in the region are the Margaret River (Beatty et al., 2007), around 1.5 times higher than precipitation comprehensive foreshore assessments (e.g. rates (Taylor & Tinley, 1999; BoM, 2015) CCG, 2005a, 2006, 2008), and research, resulting in highly seasonal flows, with monitoring and recovery work on the re- most drainage systems becoming a series of gion’s critically endangered species (e.g. isolated pools during the dry summer- Bunn, 2004; Bunn et al., 2008; Burnham et autumn period. al., 2012; Duffy et al., 2014; Morgan et al., 2014b; Klunzinger et al., 2015). Given the CATCHMENT CONDITION IN THE burgeoning amount of unpublished litera- CAPE TO CAPE REGION ture pertaining to the catchments and resi- dent aquatic macrofauna of the CCR, the Although small on a global scale, the aim of this review is to collate and interpret Margaret River is the largest river system the available information and data, thus in the CCR, draining an area of 477 km2 providing a framework for environmental and features extensive permanent pool hab- managers to develop a conservation strategy itats throughout the middle and upper that may be used to protect and enhance the reaches. The river is in relatively good con- region’s aquatic biodiversity values. dition and has not become impacted by sec- ondary salinisation, unlike many other riv- STUDY AREA ers in south-western Australia (Morgan et al., 2003; Beatty et al., 2011). Almost the This review focuses on the area be- entire upper Margaret River catchment and tween Cape Naturaliste (33.53° S, 115.00° significant portions of the lower catchment E) and Cape Leeuwin (34.38° S, 115.14° E) are forested (Green et al., 2010) and only and encompasses all intervening catch- 21% of the catchment has been cleared of ments draining westward to the Indian native vegetation (CCG, 2003; DoW, 2008). Ocean (Figure 1). The geology of the region In contrast, other CCR catchments are features a coastal strip of sandy soil overly- much smaller and most have been cleared ing limestone with occasional outcroppings more extensively (Table 1). Agriculture of the underlying granitic and gneissic bed- (predominantly livestock and viticulture) is rock (Leeuwin Block), whilst further inland the dominant land use across the CCR, with the topography is undulating and low-relief the proportion of cleared land typically ex- (20-100 m above sea level) with a mixture of ceeding 60% (Table 1). Wilyabrup Brook is gravelly and sandy soils in upland areas the most highly modified catchment in the and alluvial sandy loams on drainage lines CCR (Table 1) with unrestricted access of and floodplains (Tille & Lantzke, 1990; livestock to riparian habitats prevalent CCG, 2003). (CCG, 2006).

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FIGURE 1. Major catchments and towns of the Mediterranean climatic Cape to Cape region of Western Australia.

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TABLE 1. Attributes of major catchments of the Cape to Cape region, south-western Australia. The categorisation of foreshore condition (A – pristine; B – weedy; C – erosion prone/eroding; D - ditch) was derived from the application of methods developed by Pen and Scott (1995). Area Stream Foreshore Catchment (km2) length (km) condition Dominant land use types Key environmental issues References Yallingup Brook Native vegetation (64%); agriculture Erosion & sedimentation; degradation Taylor & Tinley, <10 5.5 Not available (36%) & destruction of riparian vegetation 1999 Erosion & sedimentation; excessive Gunyulgup A <1%; B 29%; C 58%; D Agriculture (65%); native vegetation livestock access to riparian zone; Hunt et al., 2002; Brook 47 24.3 6.5% (35%) water abstraction / barriers (110 CCG, 2005b

dams) A 6%; B 17%; C 22%; D Excessive livestock access to riparian Agriculture (mainly livestock & Wilyabrup Brook 44%; zone; intensification of land use; re- Hunt et al., 2002; 89 ~100 viticulture) (84%); native vegetation Dams 11% duced flows / water abstraction / bar- CCG, 2006 (12%); residential (4%) riers (~100 dams) Excessive livestock access to riparian Cowaramup A 15%; B 32%; C 25%; D zone; reduced flows /water abstraction Agriculture (mainly viticulture) Hunt et al., 2002; Brook 24 30 28% / barriers (19 dams); sedimentation; (64%); native vegetation (36%) CCG, 2008 salinisation of springs; limited refuge habitat Excessive livestock access to riparian A 5%; B 29%; C 27%; D Agriculture (70%); native vegetation Ellen Brook zone; expanding human population; Hunt et al., 2002; 29 40 37%; unassessed 2% (30%) water abstraction / barriers (32 dams CCG, 2005a

+ 1 weir); algal blooms Main channel: A 50%; B 45%; C 5%; D 0%. Lower Degradation & destruction of riparian Pen, 1999; CCG, tributaries: A 11%; B Forested (mostly native, some plan- vegetation; water abstraction / barrier Margaret River 2003, 2009a, 2011; 477 ~250 46%; C 20%; D 23%. tations) (79%); cleared (agriculture, (670 dams, 43 > 8 ML capacity); alien DoW, 2008; Green Bramley Brook: A 28%; residential) (21%) species introductions; pollution; live- et al., 2010 B 31%; C 6%; D 35% stock access to riparian zone

Excessive livestock access to riparian A 15%; B 38%; C 30%; D Boodjidup Brook Agriculture (49%); native vegetation zone; reduced flows /water abstraction 60 55 17% CCG, 2009b (36%); rural residential (10%) / barriers (90 dams); erosion / sedi-

mentation A 2%; B 41%; C 14%; D Water abstraction / flow reductions/ Qunninup Brook Agriculture (livestock & viticulture) Hunt et al., 2002; 19 20.4 43% barriers; degradation & destruction of (72%); native vegetation (27%) CCG, 2013 riparian vegetation

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The proliferation of on-stream dams Bostockia cf. porosa. Preliminary genetic is a key issue impacting catchments across analyses have revealed that the latter two the CCR (Table 1). Wilyabrup Brook, for taxa represent cryptic species that are as instance, has ca 100 on-stream dams (CCG, yet undescribed (Unmack, 2013; Morgan et 2006), and the density of dams is even high- al., 2014a). Additionally, five introduced er in the Gunyulgup Brook and Boodjidup teleost species have been recorded in fresh Brook catchments (Green et al., 2010; Table waters of the CCR, i.e. Rainbow Trout On- 1). The exception to this trend is the Marga- corhynchus mykiss (Walbaum 1792), Gold- ret River main channel, which has only fish Carassius auratus Linnaeus 1758, three low-level weirs, although there are Common Carp Cyprinus carpio Linneaus 670 dams on its numerous tributaries in- 1758, Eastern Gambusia Gambusia cluding 43 of commercial size (i.e. >8 ML holbrooki (Girard 1859) and Redfin Perch storage capacity) (Green et al., 2010; Table Perca fluviatilis Linnaeus 1758 (Table 2; 1). Water is also pumped directly from the Figure 3); although some of these species river in the middle and lower reaches for may no longer be present in the region (see anthropogenic uses (Green et al., 2010). A below). report on the Ecological Water Require- The CCR also houses nine decapod ments (EWR) in two representative reaches species (Table 3), seven of which of the lower and middle section of Margaret are native (Figure 4), i.e. Hairy Marron River suggested that current levels of water Cherax tenuimanus, Gilgie Cherax quinque- abstraction were within ecologically sus- carinatus Gray 1845, Restricted Gilgie tainable limits but ongoing monitoring was Cherax crassimanus Riek 1967, Koonac required to ensure these limits would not be Cherax preissii (Erichson 1846), Glossy exceeded with future allocation of water Koonac Cherax glaber Riek 1967, Margaret licenses and climate change (Green et al., River Burrowing Crayfish Engaewa pseu- 2010). In Cowaramup Brook, however, a doreducta, Augusta Burrowing Crayfish similar study found that water abstraction Engaewa similis Riek 1967, and an addi- was already close to the ecologically sus- tional two that have been introduced from tainable yield, with only a limited capacity elsewhere in Australia (Figure 3), i.e. for the system to accommodate increased Smooth Marron Cherax cainii Austin 2002 consumption (Donohue et al., 2010). and Yabby Cherax destructor Clark 1936. The freshwater bivalve mollusc Carter’s FRESHWATER MACROFAUNA OF Freshwater Mussel (Westralunio carteri) THE CAPE TO CAPE REGION also occurs in the CCR (Table 3, Figure 4). A number of euryhaline species (e.g. The CCR houses a total of six native Leptatherina wallacei (Prince, Ivantsoff & fish species (Table 2; Figure 2) including one Potter 1982), Pseudogobius olorum agnathan, the Pouched Lamprey Geotria (Sauvage 1880), Afurcagobius suppositus australis Gray 1851, and five teleosts, i.e. (Sauvage 1880)) have occasionally been re- Western Minnow Galaxias occidentalis ported from fresh waters of the CCR; how- Ogilby 1899, Western Mud Minnow Galaxi- ever, these species generally only use ella munda McDowall at1978, Balston’s freshwater habitats opportunistically and Pygmy Perch N. balstoni, Western Pygmy will not be covered further in this review. Perch Nannoperca cf. vittata and Nightfish

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FIGURE 2. Native freshwater fishes of the Cape to Cape region: a) maturing Pouched Lamprey (Geotria australis) (image: S. Beatty); b) suctorial disc of sub- adult Pouched Lamprey (image: S. Beatty); c) metamorphosed juvenile or down- stream migrant Pouched Lamprey (image: D. Morgan); d) larva (ammocoete) Pouched Lamprey (image: D. Morgan); e) Western Minnow (Galaxias occidentalis) (image: M. Allen); f) Western Mud Minnow (Galaxiella munda) (image: G. R. Al- len); g) Nightfish (Bostockia porosa) (image: S. Beatty); h) Western Pygmy Perch (Nannoperca vittata) (image: M. Allen); i) Balston’s Pygmy Perch (Nannatherina balstoni) (image: D. Morgan).

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FIGURE 3. Introduced freshwater macrofauna species of the Cape to Cape region: a) Rainbow Trout (Oncorhynchus mykiss) (image: R. Kuiter); b) Goldfish (Carassius auratus) (image: S. Beatty); c) Common Carp (Cyprinus carpio) (image: S. Beatty); d) Redfin Perch (Perca fluviatilis) (image: R. Kuiter); e) female Eastern Gambusia (Gambusia holbrooki) (image: D. Morgan); f) male Eastern Gambusia (image: M. Allen); g) Smooth Marron (Cherax cainii) (image: D. Morgan); h) Yabby (Cherax de- structor) (image: D. Morgan).

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FIGURE 4. Native freshwater decapod crustaceans and bivalve molluscs in the Cape to Cape region: a) Margaret River Burrowing Crayfish (Engaewa pseudore- ducta) (image: Q. Burnham); b) Augusta Burrowing Crayfish (Engaewa similis) (image: Q. Burnham); c) Hairy Marron (Cherax tenuimanus) (image: S. Visser); d) Gilgie (Cherax quinquecarinatus) (image: D. Morgan); e) Restricted Koonac (Cherax crassimanus) (image: S. Beatty); f) Koonac (Cherax preissii) (image: D. Morgan); g) Glossy Koonac (Cherax glaber) (image: R. McCormack); h) Carter’s Freshwater Mussel (Westralunio carteri) (image: D. Morgan).

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TABLE 2. Freshwater fish species present in the Cape to Cape region, south-western Australia. NB -* denotes an intro- duced species; ^ denotes a species that is presumed to have been locally extirpated from a catchment.

Family/Species Common name Catchments References Geotriidae Morgan et al., 1998; Morgan & Beatty, 2003, 2004a, Geotria australis Pouched Lamprey Margaret 2007a; Beatty & Morgan, 2008 Galaxiidae Morgan et al., 1998, 2013; Morgan & Beatty, 2003, Margaret, Wilyabrup, Boodjidup, Galaxias occidentalis Western Minnow 2004a, 2005, 2007a, 2008; Beatty et al., 2006, 2007, Turner 2008; Beatty & Morgan, 2008; Beatty & Allen, 2008; Morgan et al., 1998; Morgan & Beatty, 2003, 2004a, Galaxiella munda Western Mud Minnow Margaret, Wilyabrup, Boodjidup 2008; Beatty & Allen, 2008; Beatty et al., 2008 Percichthyidae Margaret, Wilyabrup, Ellen, Morgan et al., 1998, 2013; Morgan & Beatty, 2003, Bostockia porosa Nightfish Boodjidup, Turner^ 2004a, 2005, 2008; Beatty & Allen, 2008; Morgan et al., 1998, 2013; Morgan & Beatty, 2003, Margaret, Wilyabrup, Ellen, Nannoperca vittata Western Pygmy Perch 2004a, 2005; Beatty et al., 2006, 2008; Beatty & Turner^ Allen, 2008 Nannatherina balstoni Balston’s Pygmy Perch Margaret., Turner^ Morgan et al., 1998, 2013; Morgan & Beatty, 2003 Salmonidae Margaret, private farm dams *Oncorhynchus mykiss *Rainbow Trout throughout CCR Morgan et al., 1998; CCG, 2011 Cyprinidae *Carassius auratus *Goldfish Margaret^, Wilyabrup Beatty et al., 2008; Allen et al., 2013 *Cyprinus carpio *Common Carp Margaret^, Wilyabrup^ R. Paice, pers. comm.; D. McKenzie, pers. comm. Poeciliidae Morgan et al., 1998, 2013; Morgan & Beatty, 2003, Margaret, Cowaramup, Ellen, *Gambusia holbrooki *Eastern Gambusia 2004a, 2005, 2008; Beatty & Morgan, 2008; Beatty et Boodjidup, Turner al., 2008 Percidae *Perca fluviatilis *Redfin Perch Margaret^ Morgan et al., 1998

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TABLE 3. Freshwater decapod crustaceans () and bivalve mollusc (Hyriidae) species present in the Cape to Cape region, south-western Australia. NB -* denotes an introduced species; ^ denotes a species that is presumed to have been locally extirpated from a catchment; (?) denotes an unconfirmed record.

Family/Species Common name Catchments References Parastacidae Engaewa pseudoreducta Margaret River Burrowing Margaret Burnham et al., 2012; DEC 2008 Crayfish Engaewa similis Augusta Burrowing Crayfish Margaret, Boodjidup, Turner^ Horwitz & Adams 2000; Morgan et al. 2013; Q. Burnham pers. comm. Cherax tenuimanus Hairy Marron Margaret Austin & Ryan 2002; Bunn 2004; Molony et al. 2004; de Graaf et al. 2009 Cherax quinquecarina- Gilgie Margaret, Wilyabrup, Ellen, Cowa- Morgan & Beatty 2003, 2004a, 2005, 2008; Beatty et tus ramup, Gunyulgup, Boodjidup, Cal- al. 2006, 2008; Beatty & Allen 2008; gardup Cherax crassimanus Restricted Koonac Margaret, Ellen, Turner^ Morgan & Beatty 2003, 2004a, 2005, 2008; Beatty et al. 2006, 2008; Beatty & Allen 2008; Cherax preissii Koonac Margaret, Ellen, Calgardup, Turner Morgan & Beatty 2005; Morgan et al. 2013 Cherax glaber Glossy Koonac Calgardup (?) Austin & Knott 1996 *Cherax cainii *Smooth Marron Margaret, Wilyabrup, Cowaramup, Morgan & Beatty 2005, 2008; Beatty et al. 2006; de Ellen, Gunyulgup, Boodjidup Graaf et al. 2009 *Cherax destructor *Yabby Margaret, Wilyabrup, Gunyulgup, Morgan & Beatty 2005, 2008; Beatty et al. 2006, Boodjidup, Turner 2008; Beatty & Allen 2008; Allen et al. 2013; Morgan et al. 2013 Hyriidae Westralunio carteri Carter’s Freshwater Mus- Margaret, Wilyabrup, Ellen, sel Boodjidup Klunzinger et al. 2012

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THREATENING PROCESSES IN THE reserve that was established many years CAPE TO CAPE REGION prior to the initial fish survey. Whilst the spread of invasive Eastern Gambusia may be a factor in the decline of Western Mud Climate change Minnow (discussed in the Introduced spe- South-western Australia has experi- cies section below), a more likely explana- enced a 10-15% reduction in rainfall since tion for the observed fish declines in the the mid-1970s with some parts of the region Margaret River is the sharp drop in annual experiencing a 50% reduction in surface stream discharge in recent years. Five of the runoff during this time (Silberstein et al., lowest discharge years on record (since 1970 2012). Global Climatic Models unanimously when record keeping commenced in the project further reductions in rainfall (ca 8%) catchment) have occurred since 2006. West- and runoff (ca 24%), and an increase in the ern Mud Minnow has been shown to spawn number of no-flow days (by up to 4 months) in temporarily flooded riparian zones and by 2030 (Suppiah et al., 2007; CSIRO, its recruitment success is likely to be linked 2009a, b; Barron et al., 2012; Silberstein et to stream flow as has been demonstrated for al., 2012). Compared with baseline period other south-western Australian fishes in- between 1986-2005, median estimates of cluding lampreys (Morgan et al., 1998; average annual rainfall decrease for the Beatty et al., 2007, 2014). Moreover, West- region by 2090 are 12% under intermediate ern Mud Minnow typically has a short emissions scenario (RCP 4.5), and 18% un- lifespan with most individuals spawning at der high emissions scenario (RCP 8.5); the age one and perishing shortly afterwards latter could result in ca 65% reductions in (Pen et al., 1991), therefore population annual runoff (Hope et al., 2015). Drought numbers are probably susceptible to dra- periods and days of extreme heat are also matic inter-annual fluctuations depending projected to increase, which will cause high- on the suitability of environmental condi- er evaporation rates (CSIRO, 2009a, b, Hope tions from year to year. The prolonged et al., 2015). These changes are likely to drought conditions in recent years may ac- have a profound impact on freshwater eco- count for the severe population decline of systems in the region, in particular on the these species in Margaret River, as well as quantity and quality of available baseflow providing a bellwether of the potential for refuge habitat. The anticipated decline in climate change to impact freshwater species biological carrying capacity of freshwater in this system and throughout the south- ecosystems due to climate change may lead west region. to localised extirpations or even extinctions The burrowing (Engaewa of the rarest and most vulnerable species spp.) are likely to be particularly vulnerable (IPCC, 2013, 2014; DEC, 2008). to climate change as they are restricted to A recent survey of key baseflow ref- ephemeral habitats that may cease to func- uge habitats in the Margaret River has re- tion effectively in the future due to rainfall vealed dramatic declines in the abundance reductions and declining water tables of both Pouched Lamprey and Western Mud (Commander, 2000; DEC, 2008). It is un- Minnow. Pouched Lamprey larvae (i.e. am- known if these species will have the capaci- mocoetes) were not found at 75% of previous ty to burrow to sufficient depths in order to known sites and not a single individual withstand predicted water table declines in Western Mud Minnow was captured despite the region and interventionary actions may thorough sampling of sites where the spe- be required to maintain populations in their cies was previously common (see Morgan & natural habitats. The Margaret River Bur- Beatty, 2003). Land use and water quality rowing Crayfish (E. pseudoreducta) is al- in the Margaret River catchment have not ready critically endangered due to its small changed markedly over this time period; geographic range, limited dispersal capabil- indeed, the historical Western Mud Minnow ity, specialised ecological requirements and sites were all located within a conservation small population size (DEC, 2008). These 12

Threatened fauna of Australia’s Cape to Cape region 2017.002: 26 attributes combine to make it the most im- Introduced species periled aquatic species in the CCR (and ar- Over the past 40 years, the rate of guably in the entire south-western Austral- species introductions has soared (75% in- ian region), and its conservation warrants crease since 1970) in fresh waters of south- top priority. Captive breeding programs western Australia (Beatty & Morgan, 2013). may need to be considered in the future to The earliest introductions were government avoid extinction of this species (DEC, 2008). sanctioned for the purpose of creating en- hanced recreational angling opportunities Water abstraction (e.g. Redfin Perch, Rainbow Trout) and for Groundwater abstraction can reduce the biological control of mosquitos (i.e. East- flow from natural springs (CSIRO, 2009b), ern Gambusia). More recently, popular which has been shown to be vital in main- aquarium and aquaculture species (e.g. taining important aquatic refuges in some Goldfish, Carp, Yabby, Smooth Marron) catchments in south-western Australia have become established in natural water- (Beatty et al., 2014). It will be critical to ways via the dumping of unwanted ensure that abstraction from major aquifers, or via their escape from ponds and dams particularly the south-west Yarragadee and during flooding (Morgan & Beatty, 2004b, Leederville, does not exacerbate the effects 2007b; Morgan et al., 2004, 2011; Beatty & of climate change in reducing the amount Morgan, 2013). Invasive aquatic species and quality of baseflow refuge pools; partic- have been shown to have adverse impacts ularly those in the upper Margaret River. on Australian native species through preda- Other water abstraction issues include the tion, competition for food and space re- illegal pumping of river water, non- sources, agonistic behaviour, habitat degra- compliance by private land holders in the dation, and as vectors of diseases and para- installation and use of flow-bypass valves in sites (e.g. Fletcher et al., 1985; Horwitz, dams, and the impact of timber plantations 1990; Gill et al., 1999; Morgan et al., 2002, intercepting run-off and causing draw-down 2004; Morgan & Beatty, 2007b; Tay et al., of water tables (DoW, 2009). In the Cowa- 2007; Lymbery et al., 2010; Beatty & Mor- ramup Brook catchment, water abstraction gan, 2013). is already very close to the maximum sus- Six introduced aquatic macrofauna tainable level necessary to maintain aquatic species have been recorded in fresh waters ecological functions (Donohue et al., 2010). of the CCR (Tables 2; 3). Additionally, de- Further EWR studies would be beneficial in spite being a south-western Australian en- order to determine if water use is encroach- demic, translocation of Smooth Marron into ing on ecologically sustainable limits in oth- the Margaret River catchment has pushed er catchments of the region. The State Gov- the Hairy Marron to the brink of extinction ernment’s water authorities have embarked (Bunn, 2004; de Graaf et al., 2009; Duffy et on a massive public educational campaign to al., 2014). In response, the Department of raise awareness of the precarious state of Fisheries, Government of Western Austral- potable water supplies in south-western ia, has established a captive breeding pro- Australia and to encourage strategies for gram of genetically validated pure-strain limiting water wastage. Clearly, authorities Hairy Marron for supplementation of wild will need to closely monitor and, where re- stocks (de Graaf et al., 2009; Duffy et al., quired, impose restrictions on human water 2014), and has overseen the removal of use in the CCR and across the broader re- large numbers of Smooth Marron and hy- gion in order to balance human demands brid specimens from the upper Margaret with ecological requirements as the region River since 2004, which has slowed but not continues its transition to drier and warmer reversed the overall decline of Hairy Marron climatic conditions. in this system (de Graaf et al., 2009).

Generally, once established, intro- duced species prove impossible to eradicate 13

Threatened fauna of Australia’s Cape to Cape region 2017.002: 26 from the wild (Horwitz, 1990; Rowe et al., captured from this site showing signs of 2008). On the one hand, Smooth Marron caudal fin damage due to fin-nipping by and Eastern Gambusia have become well Eastern Gambusia (Figure 7). Its estab- established and widespread in the CCR lishment in Canebrake Pool may have also (Figures 5 and 6), with the latter species contributed to the decline of Western Mud recently being discovered for the first time Minnow at this site. However, as the abun- in Canebrake Pool, one of the most im- dance of Western Mud Minnow has also portant baseflow refuge habitats in the dropped to undetectable levels in refuge Margaret River (Allen et al., 2015). This pools further upstream that are not yet col- noxious pest species has rapidly become one onised by Eastern Gambusia, the decline is of the most abundant fishes at this site, un- more likely attributable to climate change derlining its highly invasive traits. Its im- driven flow reductions. pact on the native aquatic fauna has been significant, with >90% of native percicthyids

FIGURE 5. Distribution of introduced fishes in the Cape to Cape region of West- ern Australia. 14

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FIGURE 6. Distribution of intro- duced crayfishes in the Cape to Cape region of Western Australia.

FIGURE 7. Caudal fin damage on percichthyids captured from Cane- brake Pool, Margaret River (March- April 2015) caused by agonistic behav- iour of Eastern Gambusia; a) Balston’s Pygmy Perch (damaged tail above, undamaged tail below); b) Nightfish; c) Western Pygmy Perch (images: M. Allen). 15

Threatened fauna of Australia’s Cape to Cape region 2017.002: 26

The Yabby is also in the early stages tional campaign to address this growing of establishing itself in the region (Figure threat. 6). Unfortunately, this species is very adaptable to local conditions and eradica- Instream barriers tion is unlikely via conventional control Artificial barriers on waterways (e.g. methods. The Yabby is of particular concern dams, weirs, road crossings) can impact on as it occupies an equivalent ecological niche aquatic species in a number of ways. Most to endemic Cherax spp. and directly com- crucially, they reduce longitudinal connec- petes with them for food and habitat re- tivity in lotic systems, which blocks or re- sources (Beatty, 2006; Lynas et al., 2007). It stricts the access of migratory species to is also capable of breeding multiple times vital habitats such as breeding grounds, per year, allowing it to rapidly colonise hab- seasonal feeding areas, or nurseries (Lucas itats and potentially outnumber native spe- & Baras, 2001; Morgan & Beatty, 2003; cies (Beatty et al., 2005). Koehn & Crook, 2013). For example, a num- The remaining alien species have ei- ber of native freshwater fishes (e.g. Western ther failed to establish self-sustaining popu- Minnow, Nightfish) move upstream to lations (e.g. Rainbow Trout and Redfin spawn in lower order tributaries with the Perch), or their spread in the region has, onset of winter flows (Beatty et al., 2014). thus far, been effectively contained through Not only do instream barriers restrict the targeted fishing (e.g. Goldfish and Common migration of such species through physical Carp). Goldfish have been eradicated from blockage, they also delay the onset of sea- Darch Brook (a tributary of Margaret River) sonal flows that provide spawning cues, via a combination of habitat dewatering, thus shortening the duration of the breeding netting and chemical icthyocide (rotenone) season. In the CCR, hundreds of instream application (Allen et al., 2013). As far as the barriers (mainly gully dams) have been in- authors are aware, this represents the first stalled in recent decades (e.g. CCG, 2005a, successful eradication of the species from an 2006), yet only three fish passage structures aquatic system in Western Australia. The have been built during this time (Beatty et only other record of Goldfish in the CCR is al., 2007; Beatty & Allen, 2008). Further- from an artificial pond in the Wilyabrup more, these fishways have been shown to Brook catchment near Cowaramup, where effectively allow passage of only a subset of >1,000 individuals were removed in 2011 the resident fish fauna (i.e. Western Min- (Freshwater Fish Group, Murdoch Universi- now, Pouched Lamprey and Western Pygmy ty, unpubl. data). However the species re- Perch) while blocking upstream passage of mains at liberty in the catchment. other fishes (Morgan & Beatty, 2004; Beat- ty & Allen, 2008; Beatty & Morgan, 2008). The strategy of targeted fishing ef- Thus, there is a need for further research fort has had mixed success in mitigating the into modifications or alternative designs environmental risks of introduced freshwa- that will allow the full suite of migratory ter species in the CCR, but undoubtedly a native aquatic species to bypass instream more cost-effective and practical strategy barriers in the CCR and more broadly would be to prevent further introductions throughout south-western Australia. from occurring in the first instance. This is probably best achieved through educational A simpler and more effective means programs designed to raise awareness and of mitigating the impacts of instream barri- appreciation of native biodiversity whilst ers on migratory species may be the de- also emphasising the negative impacts that commissioning and removal of artificial bar- introduced species can have on native wild- riers (see Beatty et al., 2013). However, life. The biosecurity branch of the Depart- these structures are rarely redundant and ment of Fisheries (Government of Western may even provide ecological benefits such as Australia) has recently launched an educa- the provision of permanent refuge habitats in their associated impoundments (an eco- 16

Threatened fauna of Australia’s Cape to Cape region 2017.002: 26 logical function that will become increasing- native species in some of the major rivers of ly valuable in a drying climatic region), or south-western Australia (Pen, 1999; Morgan by limiting the dispersal of introduced spe- et al., 1998, 2003). cies. In Boodjidup Brook for instance, the Much riparian rehabilitation and absence of Eastern Gambusia from the ma- fencing work has already been undertaken jority of the upper catchment is presumably in the region (D. McKenzie, Cape to Cape due to the presence of an instream barrier, Catchments Group pers. comm.) and our which may, in part, explain why the West- review revealed that further work is needed. ern Mud Minnow persists in reasonable From the standpoint of conserving aquatic numbers in this system (Morgan & Beatty, fauna, future works should focus on sites 2008). that house threatened or rare aquatic spe- cies, particularly Burrowing Crayfishes Loss and degradation of riparian vege- (Engaewa spp.), Balston’s Pygmy Perch, and tation Western Mud Minnow. Riparian vegetation provides multi- ple benefits for both terrestrial and aquatic Fire fauna and plays a vital role in maintaining Native vegetation has long been function and health (Pen, burnt on a multi-annual rotational schedule 1999). In the past, riparian vegetation was as a land management tool for reducing fuel commonly destroyed during the develop- loads and associated risk of destructive wild- ment of agricultural land. The infamous fires, and to stimulate native plant growth case of the extirpation of a population of and reproduction (Christensen & Kimber, critically endangered Margaret River Bur- 1975). Given the prevalence of fire in the rowing Crayfish at the type locality in 1985 Australian landscape (and particularly in following the construction of a dam and es- the south-west of Australia); it is surprising tablishment of a timber plantation (DEC, that its impact on aquatic ecosystems has 2008) highlights the severity of the impact not yet been studied in detail. Concerns have that such practices can have on aquatic been raised about the impact of fire on macrofauna. Although the rate of clearing of Engaewa spp., especially the highly restrict- riparian vegetation has declined in recent ed E. pseudoreducta (DEC, 2008). The re- years, degradation of remnant vegetation is stricted range of this species leaves it ex- an ongoing issue across the region, mainly tremely vulnerable to habitat disturbance due to unfettered access of livestock to that may occur during fire management ac- stream lines and this threatens aquatic eco- tivities such as the clearing of firebreaks and systems through processes of erosion, sedi- containment lines, or in the use of fire re- mentation and increased nutrient inputs. tardant chemicals (DEC, 2008). In the Mature overstorey vegetation is vital Engaewa Recovery Plan it was stated that in moderating water temperature and evap- there was an “urgent need to improve the oration rates through shading (Pen, 1999; understanding of the effects of fire on Davies et al., 2007), and provides instream Engaewa spp.” (DEC, 2008) but little has habitat for native aquatic species such as been done to address this knowledge gap, Nightfish and Marron in the form of woody and we contend that the same applies to debris (Pen, 1999). Additionally, the leafy other native aquatic species, and aquatic material that falls into the water is a vital ecosystem more generally, as well. The source of carbon that drives aquatic food threat of wild fires is likely to escalate as the webs in a region where primary productivity region’s climate becomes hotter and drier; in streams is naturally low (Pen, 1999). therefore managing associated risks is likely Deep rooted riparian vegetation also keeps to become more prevalent in the future. salt-ladened groundwater at a sufficient depth to prevent secondary salinisation of streams, which has led to the extirpation of 17

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CONCLUSIONS been commendable, but our review reveals a clear need for ongoing and adaptive The Cape to Cape Region is an iconic management, as well as further monitoring part of Mediterranean climatic Australia. It and onground works that will contribute to has an expanding population and is also one the protection of the aquatic fauna values of of the premier tourist destinations in the CCR. Western Australia due to the climate and natural beauty. This review has highlighted ACKNOWLEDGEMENTS that the aquatic macrofauna of the region, whilst depauperate, has exceptional rates of endemism and contains a number of We wish to thank the Department of Parks threatened species. We recommend that and Wildlife (DpaW, formerly DEC), Gov- sites within the CCR known to house these ernment of Western Australia, for providing species (Figures 8 and 9) be monitored on a the funding to undertake this project. We regular basis in order to maintain an up-to- thank Drew McKenzie from the Cape to date knowledge of the conservation status of Cape Catchments Group for assisting with threatened populations and their habitats the project, Dr Quinton Burnham (Griffith so that immediate threats can be identified University) for providing images and valua- and addressed (e.g. riparian zone ble feedback on Engaewa, Dr Rodney Duffy degradation, water quality issues, novel (Department of Fisheries) for information species introductions). The importance of on Hairy Marron conservation initiatives, such monitoring was highlighted during the Mike Klunzinger (Murdoch University) for recent survey of baseflow refuge pools in information on freshwater bivalves, and Sue Margaret River, where a severe decline in Morrison and Andrew Hosie (Western Aus- abundance of both Western Mud Minnow tralian Museum) for contributing historical and Pouched Lamprey was detected, as was collection records. We thank Dr Gerald R. the spread of invasive Eastern Gambusia Allen (Western Australian Museum), Rudie into the upper Margaret River (Allen et al., Kuiter, Robert McCormack, and Simon 2015). Visser for allowing us to use their images. Lastly, we also acknowledge the original One of the major environmental environmental custodians and traditional challenges faced in the region is finding a owners of the CCR - the Wardandi people. balance between human water demands and those of the environment. Many of the aquatic ecosystems in the region are AUTHOR CONTRIBUTIONS degraded, while the areas that remain near pristine face a number of serious threats. Each of the authors contributed to The work on improving the management of the collection of data, and writing of the aquatic ecosystems in the CCR to date has manuscript

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FIGURE 8. Distribution of threatened fish and crayfish species in the Cape to Cape region, Western Australia. Black diamonds indicate known sites where the species is likely to remain extant; pink diamonds indicate historical sites where species is presumed to have been lost.

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FIGURE 9. Conservation priority areas in the Cape to Cape region housing threatened, endangered and priority species (i.e. Hairy Marron, Margaret River Burrowing Crayfish, Balston’s Pygmy Perch, Western Mud Minnow, and Pouched Lamprey)

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