Zoodiversity_06_2021.indb UDC 594.3(94) ON THE BIODIVERSITY HOTSPOT OF LARGE BRANCHIOPODS (CRUSTACEA, BRANCHIOPODA) IN THE CENTRAL PAROO IN SEMIARID AUSTRALIA B. V. Timms1*, M. Schwentner2, D. C. Rogers3 1Honorary Research Associate, Australian Museum, 10 William St, Sydney, 2000, and Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington, 2052, Australia 2Natural History Museum Vienna, 3rd Zoological Department; Vienna, Austria 3Kansas Biological Survey, and Th e Natural History Museum (Biodiversity Institute), Th e University of Kansas, Lawrence, KS, USA *Corresponding author E-mail: brian.timms@unsw.edu.au B. V. Timms (https://orcid.org/0000-0002-1695-148X) M. Schwentner (https://orcid.org/0000-0002-1373-456X) D. C. Rogers (http://orcid.org/0000-0003-3335-7287) On the Biodiversity Hotspot of Large Branchiopods (Crustacea, Branchiopoda) in the Central Paroo in Semi-Arid Australia. Timms, B. V., Schwentner, M., Rogers, D. C. — Biodiversity is central to the structure and functioning of communities including those of temporary water bodies. Worldwide the large branchiopod component commonly consists up to about six species instantaneously per site and twice that number across the surrounding district. Where these fi gures reach eight to ten species per site and about twice that number per district, the term diversity hotspot is sometimes used. In eastern Australia, biogeographical factors have facilitated a rich large branchiopod fauna ca 80 species and locally within 500 km2 of the central Paroo in northwestern New South Wales where a rarely diverse and abundant array of habitats supports at least 38 species, though the maximum per site syntopically is still near 10 species. K e y w o r d s : fairy shrimps, shield shrimps and clam shrimps, alpha diversity, habitat heterogeneity. Large Branchiopoda (Anostraca, Notostraca, Laevicaudata, Spinicaudata and Cyclestherida) are an important and charismatic faunal component of temporary water bodies. Although the large Branchiopoda is not a monophyletic group as are the water fl eas (Cladocera) nested within the wider Branchiopoda (Schwentner et al., 2018), they are commonly grouped together in ecological studies due to their overall life history similarities. Reports abound on alpha diversity (species richness) of the large branchiopod fauna of the world (e. g. Brendonck et al., 2008; Rogers, 2009), of continents (e. g. Anostraca of North America, Rogers, 2014; Branchiopods of South America, Rogers et al., 2021; Spinicaudata of Australia; Schwentner et al., 2015 a), large parts of continents (e. g. Anostraca in Southern Africa, Hamer and Brendonck, 1997), large countries (e. g. Large branchiopods of India, Rogers and Padhye, 2015), smaller countries (e. g. Anostraca of Botswana, Brendonck and Riddoch, 1997), districts (e. g. large branchiopods of the Chaouia Plain of western Morocco, Th iery, 1991), small groups of wetlands (e. g. Galapagos Islands, Brendonck et al., 1990) and even single wetlands (e.g. Hamer and Appleton, 1991; Wang et al., 2010). Such reports may not necessarily be complete due to further discoveries or taxonomic revisions aft er their publication, but are essential in comparative diversity studies of regional or global patterns of diversity. In some cases, there appears to be a particularly high number of the species per unit area in which case the term ‘hotspot’ may be used. Th is has been applied continentally (e. g. Spinicaudata in Zoodiversity, 55(6): 439–450, 2021 DOI 10.15407/zoo2021.06.439 Fauna and Systematics 440 B. V. Timms, M. Schwentner, D. C. Rogers Australia, Schwentner et al., 2015 a), by country or district and even to individual sites (e. g. Bovin et al, 2018 for a district; Weise, 1964 for a single site). Th ere is no rule as to how to apply this term, e. g. no defi ned number of species per unit of area, and indeed in many cases use of the term appears to be overrated when compared to diversity in other areas. In the Australian context, the region with the most large branchiopod species is the Paroo catchment (area 73,600 km2) in northwestern New South Wales (NSW) (Timms and Sanders, 2002; Timms and Richter 2002; Schwentner et al., 2015 a). Th is can be narrowed down to a much smaller segment without much loss of diversity to about 500 km2 in the central parts of the Paroo catchment. Here we detail this fauna, to consider why it should be so diverse and to consider how best to defi ne a biodiversity ‘hotspot’. Material and methods Th e area of study is three privately owned adjacent station properties, Bloodwood, Tredega and Muella in the central Paroo, 130 km northwest of Bourke in northwestern New South Wales (NSW) (fi g. 1). Th ese stations (combined area about 500 km2) are vegetated with semiarid scrub on a dune fi eld on the southern and western half and subdued ridges of brown earths on the eastern half (Hazelton and Johnson, 1973). Th e overall landscape contains many temporary lakes plus other depressions of various sizes, depth, and hydroperiod. Following signifi cant rainfall some to all of these partially or completely fi ll to form a variety of seasonally astatic aquatic habitats (table 1; fi g. 2) characterized by diff ering salinity, turbidity and hydroperiod as well as sizes, pH and the presence/absence and type of submerged vegetation. Th ese dryland wetlands are a signifi cant feature of the Paroo catchment and adjacent Bulloo and Warrego systems (combined area 210,000 km2), with some areas especially rich in large lakes and claypans as in the Currawinya National Park, a Ramsar site (Timms, 1999; 2008). While wetlands may be densest at Currawinya, a limited area of ca 500 km2 on Bloodwood, Tredega and Muella in NSW arguably has greatest variety of wetlands within these semi-arid catchments. Following signifi cant rainfall events, many wetlands on these stations were sampled for large branchiopods with D-shaped pond nets with mesh sizes 1 to 2 mm. Usually the net was swept randomly through littoral areas, but in some sites notably Gidgee Lake, the fairy shrimp, Branchinella buchananensis actively avoided nets, so a search and quick snatch method was needed. Generally branchiopod populations reached a maximum 1–2 months aft er wetland fi lling, so that collections taken within this period were used for assessment of species richness. However in some wetlands, particularly in grassy pools and some other small sites, population development was quicker so that signature collections were made two to four weeks aft er site fi lling. Collections made earlier than these windows were full of immature specimens diffi cult to identify, or if later, missing some quickly developing species. One frustrating aspect of collecting was access aft er rainfall events. Th is was partly due to station tracks being impassable sometimes for weeks, but also contributed to by public unsealed roads being closed aft er rains. Samples were taken from the wetlands over a 35-year period, 1987–2021 inclusive. However not all types of wetlands were studied simultaneously, so that there was specialization on various wetland types over the years (see table 1). Some wetlands were studied more intensely, which may have biased the data, and of course Fig. 1. Map of Bloodwood, Tregeda, and Muella Stations, central Paroo, northwestern NSW. Code to symbols: SL — Salt Lake; L — Freshwater Lake; C — Claypan; G — Grassy swamp; S — Samphire swamp; X — Poplar Box fl at; short line, creek pool; dot — Black Box swamp. 441On the Biodiversity Hotspot of Large Branchiopods in Semi-Arid Australia there were diff ering numbers of each wetland type as dictated by the landscape. So no claim is made for the exact internal comparability of the data. However, the long study period is suffi cient to encounter all extant species in the 500 km2 study area and for common combinations of species instantaneously present to be recorded at least for the most numerous wetland types. Species were identifi ed using early working versions of Timms (2012, 2015 b, 2018) with representatives of the major groups illustrated in fi g 3. In addition molecular analyses was conducted once for many species of Eulimnadia (Schwentner et al., 2015 a) Eocyzicus (Schwentner et al., 2014; Tippelt and Schwentner, 2018), Ozestheria (Schwentner et al., 2015 b; Schwentner et al., 2020) and Triops (Meusel and Schwentner, 2017). In our count of species, we distinguish between formally described species and species known and delimited only genetically. Of the latter, we only included those species that were genetically unambiguously delimited from all other species of the respective genus. In a few cases, additional rather divergent lineages whose status is questionable were identifi ed within some species, (e. g., Ozestheria sp. Q1–Q5; Schwentner et al., 2020); these were not included in the species count so as not to artifi cially infl ate the observed diversity. Th ree commonly encountered species — Triops australiensis, Ozestheria packardi and Ozestheria berneyi — were genetically shown recently to comprise more than one species each (Schwentner et al., 2015 b; Schwentner et al., 2020; Meusel and Schwentner, 2017). Although we attempted to treat each of these species separately whenever possible, for some summary statistics (e. g. species co-occurrences) we had to lump them each under one taxon, as they cannot be retrospectively separated. Because it is likely that the “true” Triops australiensis and Ozestheria packardi do not occur in the middle Paroo, but morphologically similar species do, we have counted these two species as “undescribed”. Fig. 2. Images of seven types of wetlands in the central Paroo., northwestern New South Wales: A — Gidgee Salt Lake; B — Ski Freshwater Lake; C — Melaleuca claypan; D — Beverley’s grassy pool (dry); E — Reedy Black Box swamp; F — Utah Poplar Box fl at; G — Lower Crescent creek pool. Not to scale. 442 B. V. Timms, M. Schwentner, D. C. Rogers Results Th e species encountered are listed in tables 2 to 5, together with data on their relative abundance and habitat preferences. Th e total is 38 species. Th is includes 16 species of Anostraca, dominated by 13 species of Branchinella, one species each of Laevicaudata and of Cyclestherida, and 20 species of Spinicaudata (in the seven spinicudatan genera there are multiple species of Eocyzicus (six) and Limnadopsis (fi ve)) (tables 2, 3, 4). However, a number of still undescribed species greatly increase the number of species recorded for Spinicaudata (total of 33 described and undescribed species) and Triops (total of six undescribed species) (table 7). Most of the undescribed Spinicaudata are Ozestheria species (Schwentner et al., T a b l e 1 . Wetland Types in the central Paroo, northwestern New South Wales Wetland No. Hydrology Turbidity Conductivity Reference Treed Swamps ca 50 Two physical types. Black Box Swamps fi ll every 2–3 years, Poplar Box Flats every 3–5 years. Persist 3–6 mths. 50–300 NTU 50–500 mS/cm Timms, in press Claypans ca 40 Fill every 2–3 years, more in La Nina years. Persist 3–6 months, more in La Nina years. 1000–5000 NTU 100–1000 mS/ cm Hancock and Timms, 2002 Grassy pools/ swamps 5 Fill rarely, mainly in strong La Nina years. Persist only a few weeks. < 50 NTU < 100 mS/cm Timms, 1997 Creek Pools 8 Fill most years, sometimes 2–3 times. Persist 3–36 mths. 50–500 NTU 100–10000 mS/ cm Timms, 2001 Salt Lakes 6 Of various salinity ranges. Persist 3–30 mths, mainly in La Nina years. Oft en dry for years in El Nino conditions. 10–100 NTU 1000–250000 mS/cm Timms, 1993,1998, 2018 Samphire Swamps 5 Fill every 3–5 years, certainly in La Nina years. Persist 3–6 months. < 100 NTU 200–5000 mS/ cm Schwentner et al .,2012 Freshwater Lakes 5 Episodic, lasting 6–24 mths, Ski L fi lls every 2–3 yrs, Wirrania every 2–4 years, L. Muella every 5–8 yrs Ski 200–500 NTU, others 20–100 NTU All 100–750 mS/ cm Timms, 1997 Table drains many Occasionally support branchio- pods. Persist only a few weeks in very wet conditions. < 100 NTU < 100 mS/cm Farm dams many Most are permanent and sunk into Black Box Swamps. Some dug out on slopes. These persist for a few mths. turbid fresh N o t e. NTU — Nephrometric Turbidity Unit. Fig. 3. Four representative branchiopods from the central Paroo. A — Anostracan Branchinella australiensis; B — Notostracan Triops sp.; C — Spinicaudatan Limnadopsis tatei; D — Spinicaudatan Ozestheria lutraria. 443On the Biodiversity Hotspot of Large Branchiopods in Semi-Arid Australia T a b l e 2 . Fairy shrimps (Anostraca) of the central Paroo, northwestern New South Wales Species Spread in central Paroo Distribution beyond central Paroo Regularity of appearance Habitat range Australobranchipus parooensis Rogers et al., 2007 < 5 sites SW Qld and West NSW Rare Fresh waters, mainly clear Branchinella affi nis Linder, 1941 > 25 sites Australia wide Most fi llings Fresh waters, mainly inter- mediate clarity Branchinella angelica Timms, 2016 1 site Western NSW Rare Fresh waters, mainly inter- mediate clarity Branchinella arborea Ged- des, 1981 > 50 sites Central eastern inland All fi llings Fresh waters, mainly clear Branchinella australiensis (Richters, 1876) > 50 sites Australia wide All fi llings Fresh waters, mainly inter- mediate clarity Branchinella buchananensis Geddes, 1981 < 5 sites North and central eastern inland Some fi llings Hypo and mesosaline waters Branchinella budjiti Timms, 2001 > 50 sites Central eastern inland Most fi llings Fresh waters, mainly turbid Branchinella campbelli Timms, 2001 < 5 sites Central eastern inland Some fi llings Clear fresh waters, Branchinella frondosa Henry, 1924 < 10 sites North and central Aust. Some fi llings Freshwaters, mainly of intermediate clarity Branchinellla lyrifera Linder, 1941 > 50 sites Australia wide All fi llings Turbid to extremely turbid waters Branchinella occidentalis Dakin, 1914 < 50 sites Australia wide All fi llings Turbid to extremely turbid waters Branchinella pinnata Ged- des, 1981 < 25 sites North and central Aust. Most fi llings Fresh waters, mainly inter- mediate clarity Branchinella proboscida Henry, 1924 < 50 sites Australia wide Most fi llings Mainly turbid waters Branchinella wellardi Mil- ner, 1929 < 5 sites North and central Aust. Some fi llings Clear fresh waters Parartemia minuta Geddes, 1973 < 5 sites Eastern inland Most fi llings Saline waters Streptocephalus archeri Sars, 1896 < 5 sites Australia wide Some fi llings Fresh waters, mainly inter- mediate clarity T a b l e 3 . Shield shrimps (Notostraca) of the central Paroo, northwestern New South Wales* Species Wetland types with records Distribution beyond middle Paroo Examples Triops sp. A Black Box swamps within 500 km Marsilea Pan Triops sp. B Black Box swamps, Poplar Box fl ats, Claypans within 1000 km Marsilea Pan, Turkey Claypan Triops sp. I Creek pools within 250 km Lower Crescent pool Triops sp. L Freshwater lake Australia wide Lower Lake Eliza Triops sp. N Black Box swamps, Poplar Box fl ats within 1000 km Carols Poplar Box fl at Triops sp. O Poplar Box fl ats, Creek pools, Samphires Within 500 km Carols Poplar Box fl at Lower Crescent pool Roszkos Samphire *Data from Meusel and Schwentner, 2017 using their notation. 2015 b). Syntopic occurrences are common within Branchinella, Eulimnadia, Limnadopsis, Eocyzicus, Ozestheria and Triops. Th e grand total is at least 56 species of large branchiopods within the study area. Not all of the wetlands within the central Paroo were equally speciose (table 5). High diversity in Black Box swamps and claypans is well supported with many sites studied and many samples (tables 1, 5). Poplar Box fl ats, another variety of treed swamps unique to inland eastern Australia, are also speciose, despite their generally shorter hydroperiods 444 B. V. Timms, M. Schwentner, D. C. Rogers T ab le 4 . L is t o f c la m sh ri m ps (L ae vi ca ud at a, S pi ni ca ud at a, C yc le st he ri da ) r ec or de d in th e ce nt ra l P ar oo , n or th w es te rn N ew S ou th W al es Sp ec ie s Sp re ad in ce nt ra l Pa ro o D is tr ib ut io n be yo nd c en - tr al P ar oo R eg ul ar ity o f a p- pe ar an ce H ab ita t r an ge Ly nc eu s m ac le ay an us (K in g 18 55 ) < 5 si te s A us tr al ia w id e So m e fi l lin gs C le ar fr es h w at er s, 3– 12 m th h yd ro pe ri od A us tr al im na di a gr ob be ni D ad ay , 1 92 5 1 si te o nl y N or th er n A us tr al ia R ar e C le ar fr es h w at er s, 3– 12 m th h yd ro pe ri od Eu lim na di a au st ra lie ns is T im m s, 20 16 < 5 si te s Q ld a nd N SW So m e fi l lin gs Fr es h w at er s o f < 3 m th h yd ro pe ri od Eu lim na di a be ve rl ey ae T im m s, 20 16 < 5 si te s En de m ic ? So m e fi l lin gs C le ar fr es h w at er s, 1– 2 m th h yd ro pe ri od Eu lim na di a ca na lis T im m s, 20 16 < 10 si te s En de m ic ? So m e fi l lin gs Fr es h w at er s o f < 3 m th h yd ro pe ri od Eu lim na di a ha ns on i T im m s, 20 16 < 10 si te s W ith in 2 50 k m M os t fi ll in gs Fr es h w at er s o f < 3 m th h yd ro pe ri od Li m na do ps is bi rc hi i ( Ba ir d, 1 86 0) > 10 0 si te s In la nd A us tr al ia A ll fi l lin gs A ll ty pe s o f f re sh w at er s Li m na do ps is bl oo dw oo de ns is Sc hw en tn er , T im m s a nd R ic ht er , 2 01 2 3 si te s o nl y En de m ic ? So m e fi l lin gs Sa m ph ir e sw am ps , f re sh to h yp os al in e Li m na do ps is pa ra ta te i S ch w en tn er , T im m s a nd R ic ht er , 2 01 2 < 5 si te s W ith in 2 50 k m So m e fi l lin gs V ar ie ty o f f re sh w at er s Li m na do ps is pa rv isp in us H en ry , 1 92 4 > 50 si te s In la nd Q ld a nd N SW M os t fi ll in gs M an y fr es h w at er s, es p. B la ck B ox S w am ps Li m na do ps is ta te i S pe nc er a nd H al l, 18 96 > 50 si te s In la nd A us tr al ia M os t fi ll in gs A ll ty pe s o f f re sh w at er s Pa ra lim na di a qu ee ns la nd ic us T im m s, 20 16 > 20 si te s In la nd Q ld a nd N SW M os t fi ll in gs Fr es h w at er s o f < 3 m th h yd ro pe ri od Eo cy zi cu s a rg ill aq uu s T im m s a nd R ic ht er , 2 00 9 < 10 si te s In la nd Q ld , N SW , a dj . S A M os t fi ll in gs M ai nl y tu rb id c la y pa ns Eo cy zi cu s a rm at us T ip pe lt an d Sc hw en tn er , 2 01 8 < 5 si te s N or th er n an d ce nt ra l A us tra lia M an y fi l lin gs C le ar fr es h an d hy po sa lin e w at er s Eo cy zi cu s p ar oo en sis R ic ht er a nd T im m s, 20 05 < 10 si te s En de m ic ? A ll fi l lin gs H yp os al in e st ag es o f s al t l ak es Eo cy zi cu s p hy to ph ill us T ip pe lt an d Sc hw en tn er , 2 01 8 < 5 si te s W ith in 5 00 k m M os t fi ll in gs V eg et at ed fr es h w at er s, 3– 12 m th h yd ro pe ri od Eo cy zi cu s r ic ht er i T ip pe lt an d Sc hw en tn er , 2 01 8 < 10 si te s W ith in 1 00 0 km M os t fi ll in gs Tu rb id w at er s, in c. cla yp an s a nd ca ne gr as s s w am ps Eo cy zi cu s u bi gu us T ip pe lt an d Sc hw en tn er , 2 01 8 < 10 si te s W id es pr ea d M os t fi ll in gs A ny fr es hw at er si te , b ut ra re ly tu rb id w at er s O ze st he ri a lu tr ar ia (B ra dy , 1 88 6) < 10 si te s W ith in 1 00 0 km So m e fi l lin gs M an y w et la nd ty pe s O ze st he ri a ru br a (H en ry , 1 92 4) 1 si te o nl y w ith in 1 00 0 km F re q. u nk ow n C re ek p oo l O ze st he ri a sp . A 2 < 5 si te s w ith in 1 00 0 km Fr eq u nk no w n M an y w et la nd ty pe s O ze st he ri a sp . B 2 1 si te o nl y w ith in 1 00 0 km ra re Bl ac k Bo x sw am p O ze st he ri a sp . I 2 < 5 si te s en de m ic in P ar oo Fr eq . u nk no w n M ai nl y Bl ac k Bo x sw am ps , a ls o tu rb id w et la nd s O ze st he ri a sp . K 2 1 si te o nl y w ith in 5 00 k m R ar e Ba ck B ox sw am p O ze st he ri a sp . M 1 > 10 si te s w ith in 1 00 0 km Fr eq . u nk no w n M an y w et la nd ty pe s O ze st he ri a sp . N I < 5 si te s w ith in 1 00 0 km Fr eq . u nk no w n M an y w et la nd ty pe s O ze st he ri a sp . Q 2 < 10 si te s A us tr al ia w id e Fr eq M an y w et la nd ty pe s O ze st he ri a sp . S 2 < 10 si te s w ith in a 1 00 0 km un kn ow n Po pl ar B ox fl at s Eo le pt es th er ia n r. tic in en sis B al sa m o- C ri ve lli , 1 85 9 1 si te o nl y ra re Bl ac k Bo x sw am p C yc le st he ri a nr . h isl op ii, B ai rd , 1 85 9 1 si te o nl y N or th er n A us tr al ia ra re Po pl ar B ox fl at 1 S um m ar iz ed p re vi ou sl y un de r n am e O . b er ne yi (G ur ne y, 1 92 7) ; 2 su m m ar iz ed p re vi ou sl y un de r t he n am e O . p ac ka rd i ( Br ad y, 1 88 6) . 445On the Biodiversity Hotspot of Large Branchiopods in Semi-Arid Australia (Timms, in press). Th e remaining four wetland types are based on few sites, though generally at least 20 samples were taken over the years per site. Th ey are less speciose, certainly the salt lakes, though the situation in creek pools and grassy pools is not as clear. Th e two creek pools included in this study are the most speciose of the eight initially studied (Timms, 2001) but it is hard to characterize them by a standard aggregation of species as composition is variable as indicated by a total species list of eight species each for both fairy and clam shrimps. Th ese two sites are regularly fl ushed and though they may have some resident species, many are probably introduced from overfl ow from other habitat types depending on rainfall distribution. It is possible the data for grassy pools are incomplete as few sites were available for study and they fi lled so few times. However, their lower number of anostracans then elsewhere may be valid due to the apparent adverse infl uence of thick vegetation on anostracan life styles. Some species are habitat specialists, others more catholic. While Parartemia minuta is recorded as a salt lake specialist, it is found in only two of the fi ve salt lakes studied, but there in large numbers. Th e clam shrimp Eocyzicus parooensis is more widely distributed in the salt lakes (four sites out of fi ve), but it appears only in the initial brief freshwater and T a b l e 5 . Comparison of diversity of branchiopods in the wetlands of central Paroo, northwestern NSW Wetland Type No. of sites No of samp- les Years of study No. of fairy shrimp sp. Dominant species No. of clam shrimp sp. Dominant species Average no. of sp. per sample Black Box swamps 22 117 Mainly 2010–2021 6 B. arborea B. australiensis 8 L. birchii L. parvispinus O. packardi 6.41 Poplar Box fl ats 6 15 Mainly 2010–2021 6 B. arborea B. campbelli 7 L. parvispinus L. queenslandicus O. packardi 5.86 Claypans 15 55 1998; 2010–2021 7 B. affi nis B. lyrifera B. occidentalis 4 E. argillaquus O. lutraria O. packardi 6.27 Lakes 3 33 mainly 2010–2021 5 B. australiensis B. arborea 5 O. lutraria O. packardi 3.75 Creek Pools 2 20 1998; 2010–2021 8 B. australiensis 8 L. birchii O. packardi 4.05 Grassy Pools 3 12 1987–2012 4 none 8 Eulimnadia spp. Limnadopsis spp. 4.58 Salt Lakes 5 33 1987– 2021 3 P. minuta 3 E. parooensis 1.09 Ta b l e 6 . Number of co-occurences of branchiopods in diff erent types of wetlands No. of species Black Box Swamps Poplar Box Flats Claypans Freshwater Lakes Creek pools Grassy Pools Salt Lakes 0 0 0 0 0 0 0 9 1 0 0 0 0 0 0 14 2 1 0 0 6 3 2 8 3 2 1 0 8 3 4 2 4 6 1 3 10 6 3 5 21 4 8 6 6 3 6 48 4 21 3 2 2 7 20 3 17 8 12 2 6 9 5 10 2 446 B. V. Timms, M. Schwentner, D. C. Rogers subsaline stages. Limnadopsis bloodwoodensis is found only in the freshwater and subsaline early stages of samphire swamps. Eocyzicus argillaquus is characteristically found only in quite turbid waters, hence its dominance in many claypans. Eulimnadia spp occurs in sites with short hydroperiods and is rarely found beyond the grassy pools and Poplar Box fl ats. Paralimnadia is more common and widespread, but this may be an artifact due to its wider habitat preferences and longer life cycle. Among the anostracans, Branchinella lyrifera and B. occidentalis arecharacteristic of the turbid claypans, but occasionally are found also in other moderately turbid sites. Branchinella campbelli is usually restricted to the clear waters of Poplar Box fl ats and B. wellardi to all clear waters. Limnadopsis birchii is also catholic, appearing in almost all sites in major fi lls, including briefl y in the freshwater and subsaline stages of some salt lakes! Normally it is a characteristic of Black Box swamps. Limnadopsis parvispinus is characteristic of the two types of treed swamps, so given the abundance of treed swamps, it too was common. Th e most common anostracans are Branchinella australiensis and B. arborea. Both are associated with mildly turbid waters, a common condition in Black Box swamps, freshwater lakes, creek pools, and claypans at least on initial fi lling. Discussion Fift y-six species — of which thirty-eight are formally described — are known from Bloodwood, Tredega and Muella Stations and were all found in an area of nearly 500 km2. If the southwestern half of Bloodwood Station (ca 280 km2) with by far the most wetlands is considered alone the list is still 34 described species (including all 16 anostracans). Timms and Sanders (2002) and Timms and Richter (2002) originally studied an area of 2000 km2 and counted 29 species of large branchiopods at that time. Many species were overlooked due to limited taxonomic understanding at that time. If the whole of Queensland and the adjacent northern quarter of NSW from 32o N (area 2.02 x 106 km2) which includes the Paroo study area are included, the list increases to 83 species of which 52 are formally described (table 7) with the proviso there are areas within this vast domain not yet sampled. Variations in diversity between continents (Brendonck et al., 2008; Rogers, 2009, 2014 a, b, 2015; Rogers and Timms, 2014) usually have complex contributing factors including biogeographical bases and are beyond detailed contemplation here. It is those for local T a b l e 7 . Large branchiopods in the central Paroo (Bloodwood, Tredega, Muella Stations) and also in Queensland and far northern New South Wales Taxonomic group Paroo species + undescribed All Qld + Nth NSW (N of 32oS) + undescribed Branchinella 13 – 17 – Australobranchipus 1 – 2 – Streptocephalus 1 – 1 – Parartemia 1 – 1 – Lynceus 1 – 2 1 Eulimnadia 4 4 5 9 Paralimnadia 1 – 5 – Limnadopsis 5 – 6 – Australimnadia 1 – 1 – Eoleptestheria 1 – 1 – Eocyzicus 6 – 8 – Ozestheria 2 8 2 10 Cyclestheria 1 – 1 1 Triops 0 6 0 10 Total 38 18 52 31 N o t e . Based on Meusel and Schwentner, 2016; Schwentner, 2020; Schwentner et al., 2009, 2012 a, b, 2013, 2014, 2015; Timms, 2009, 2015; Timms & Schwentner, 2012. 447On the Biodiversity Hotspot of Large Branchiopods in Semi-Arid Australia areas (say less than about 1000 km2) where marked discrepancies in diversity can occur and which oft en can at least be partly explained by various ecological factors, including climate, habitat characteristics and biotic factors (Hamer and Appleton, 1991; Rogers, 2014 a, b; Rogers and Timms, 2014; Petrov and Cvetković, 1997). Some of the more speciose examples are listed in table 8. Of these about 14 species in 500 to 1000 km2 seems close to a maximum diversity and generally contributed to by 50 % anostracans, 7 or 14 % notostracans and the remainder spinicaudatans (table 8) and sometimes a laevicaudatan (Roessler, 1995). Instantaneous coexistences (syntopy) for individual pools average less than fi ve (table 7 and table 4 in Nhiwatiwa et al., 2014) but can reach 10, and rarely, 12 species. While sites with these numbers could be considered hotspots, it is mainly Nhiwatiwa et al. (2014) who consider their data from Save Conservancy, Zimbabwe indicate a hotspot of diversity with its list of 16 species and maximum syntopy of 12 species. However, all these implied and outspoken claims fade into insignifi cance compared with the diversity in the central Paroo, Australia (tables 2, 3, 4, 7) with its 56 species or ~10 % of the globally known diversity of large Branchiopoda in this comparably small region. Surely this is a “super” hotspot of large branchiopod diversity. Interestingly the maximum instantaneous syntopy of species is still in the same order of magnitude as other hotspot contenders, i. e. about 10 to 12 species. Th is fi gure may be the maximum possible for syntopic species given their various separable niches (Th iery, 1991; Nhiwatiwa et al., 2014). Th e high species diversity in the middle Paroo is based on its high habitat diversity, with at last seven distinct types of wetlands all within close proximity (tables 1, 5; fi gs. 1, 2). Th ese have diff erent ranges of hydroperiod, turbidity, salinity, and habitat complexity which infl uence diversity of the large branchiopod inhabitants (Timms and Sanders, 2002; Timms and Richter, 2002). Most other speciose groups of sites explain their diversity on diff erences in areas, depths, and hydroperiod (e. g. Hamer and Appleton, 1991; Petrov and Cvetković, 1997). Changes during a seasonal fi lling (e. g. Melanic ponds in Serbia, Petrov and Cvetković, 1997) are of little importance in the Paroo, though two instances occur regularly there. In the saline lakes there is early colonization by Eocyzicus parooensis followed by Parartemia minuta in the hypersaline lakes and by Branchinella buchananensis in the mesosaline Gidgee Lake. Also in the claypans, predation by Branchinella occidentalis on other anostracans could infl uence the proportions of species present during a fi lling, though this has not been specifi cally detailed (Rogers and Timms, 2017; Hancock and Timms, 2002). T a b l e 8 . Large Branchiopod species richness in areas about the same size as the central Paroo Studied area Areakm2 No. of samples No. of sp. Anostraca No. of sp. Notostraca No. of sp. clam shrimps Maximum syntopy Reference Greater Bloodwood Australia 440 117 16 8 40 10 Th is study Save Conservancy Zimbabwe 150 36 8 1 7 12 Nhiwatiwa et al., 2014. Chaouia Plain Morocco 550 59 7 2 2 10 Th iéry, 1991 Kiskunság NP Hungary 500 89 6 2 3 5 Boven et al., 2008 Morava Floodplain Austria and Slo- vakia 500 – 3 2 5 – Schernhammer, 2020 and pers. com. Part of KwaZulu- Natal, South Africa 1100 10 7 1 6 10 Hamer and Appleton, 1991 Doñona NP Spain 1200 – 6 1 2 4 Diaz-Paniagua et al., 2010 Melenic, Banat prov. Serbia 0.003 – 5 2 3 7 Petrov and Cvetković, 1997 448 B. V. Timms, M. Schwentner, D. C. Rogers Sometimes diversity is enriched by the area being the junction of biogeographic provinces as it may be in the Zimbabwe example (Nhiwatiwa et al., 2014). Th is is not the case in the Paroo, but the rich branchiopod fauna across Australia provides a larger than normal pool of species that contributes to Paroo’s elevated diversity (Rogers and Timms, 2014; Schwentner et al., 2015 a). Four species are apparently endemic to the central Paroo, Eulimnadia beverleyae, E. canalis, Eocyzicus parooensis and Limnadopsis bloodwoodensis while a few more like Branchinella angelica, Eulimnadia hansoni and Limnadopsis paratatei and have limited distributions beyond the Paroo (tables 2, 4). By far the majority of species have a wide distribution, sometimes extending to central or even Western Australia. It is likely that the extensive diversity we observed in the central Paroo has not necessarily evolved in this small area, but that it is an amalgam of species that colonized this area over a long period of time (Rogers and Timms, 2014). Th eir ability to successfully colonize the central Paroo catchment was probably facilitated by its rich diversity of habitat types. It remains to consider just what constitutes a hotspot in large Branchiopod diversity. Th e situation in the Paroo is exceptional and to consider it as the only hotspot worldwide in large branchiopod diversity detracts from other diverse places. In many countries the number of branchiopod species may exceed the fi gures dealt with here, but their area is much, much larger. Two examples will suffi ce: India has about 22 species of anostracans found over 3.3 million km2 (Padhye et al., 2017) and southern Africa (i. e. the area south of the Kunene, Okavango and Zambezi rivers) has 46 species of large branchiopods over an area of 3.8 million km2 (Hamer and Brendonck, 1997). Whole biogeographic regions not surprisingly have many species of all branchiopod groups, e. g. about 200 in the Nearctic (Brendonck et al., 2008; Rogers, 2009). Th ere has been no suggestion that these large areas are termed hotspots, no matter what their diversity or specifi c area. Instead the focus is on smaller areas or individual sites, more or less about the size considered here, i. e. up to about 1000 km2. For these, a fauna of more than about 14 species and a maximum instantaneous coexistence of 8–10 or more species, as listed in table 7 and also in table 4 in Nhiwatiwa et al., 2014 is unusual and herewith is considered a hotspot for diversity. Such an acceptance would make the Paroo situation a super hotspot! Conclusions A 35 year study of 56 sites of seven types of wetlands (table 1) within a 500 km2 part of the central Paroo in northwestern NSW yielded 38 described species and a further 18 undescribed (but molecularly defi ned) species of large branchiopods (table 7). Th ese fi gures are unprecedented elsewhere in areas of similar size. Many species occur characteristically in each hydroperiod but some appeared only rarely. Instantaneous species per wetland type averaged from 6.41 in Black Box swamps to 1.09 in salt lakes (table 5). Maximal syntopic species ranged from 10 in Black Box swamps, eight in Polar Box fl ats and claypans, six in freshwater lakes, creek pools and grassy pools and just three in salt lakes (table 6). While a rich continental fauna contributed to these fi gures, the main reason is the variable seven distinct wetland types diff erentiated by salinity, turbidity, hydroperiod and habitat complexity so that the component species select distinctive habitats most suited to their ecological needs. Elsewhere in the world it is common to record up to about six syntopic species per wetland and about twice that number in the wider district (defi ned here at about 500 km2). Some wetland districts record eight-12 syntopic species, and < 20 species district wise (table 8), this apparently being the upper limit imposed by non-overlapping niche requirements. Such situations are unusual and perhaps termed hotspots. If so, then the Paroo with its extreme alpha diversity is a super hotspot. BVT is indebted to the Hansons of Bloodwood and the Batys of Muella and Tredega for access to their properties and for various other kind logistic help over the years. MS and DCR also visited the area and were most appreciative of this hospitality. 449On the Biodiversity Hotspot of Large Branchiopods in Semi-Arid Australia References Bovin, L., Vanschoenwinkel, B., De Roeck, E. R., Hulsmans, A., Brendonck, L. 2008. Diversity and distribution of large branchiopods in Kiskunság (Hungary) in relation to local habitat and spatial factors: implications for their conservation. Marine and Freshwater Research, 59, 940–950. Brendonck, L., Rogers, D. C., Olesen, J., Weeks, S., Hoeh, W. R 2008. Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater. Hydrobiologia, 595, 167–176. Brendonck, L., Riddoch, B. 1997. Anostracans (Branchiopoda) of Botswana: Morphology, distribution, diversity and endemicity. Journal of Crustacean Biology, 17 (1), 111–134. Brendonck, L., Th iery, A., Coomans, A. 1990. Taxonomy and biogeography of the Galapagos Branchiopod fauna (Anostraca, Notostraca, Spinicaudata). 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Received 21 September 2021 Accepted 3 November 2021 << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /DetectCurves 0.0000 /ColorConversionStrategy /CMYK /DoThumbnails false /EmbedAllFonts true /EmbedOpenType false /ParseICCProfilesInComments true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments true /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /CreateJDFFile false /Description << /ARA /BGR /CHS /CHT /CZE /DAN /DEU /ESP /ETI /FRA /GRE /HEB /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. 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