VERTEBRATE FAUNA N.L. McKenzie, lK. Rolfe and W.K. Youngson

Background The philosophies underlying our approach to this vertebrate survey, the sampling strategy adopted, and the actual methods we employed are described in Biological Surveys Committee ofWestern Australia (1984). Specimens representing the array of reptile and mammal species encountered in the Sandstone-Sir Samuel and Laverton-Leonora Study Areas during our survey have been lodged in the Western Australian Museum within the following series of register numbers: R62735­ 62872, 70827-70877, 78545-78585, 79000, R114956-1149570, 114983-114985; M20395, 20396, 20404-20409, 20420, 20421, 20424, 20425, 20427, 20428, 20446-20448, 20461­ 20463, 20815, 20816, 20821-20823, 20825-20830, 20832-20834, 20837, 20838, 20840,

Table 2 Faunal survey of the Sandstone-Sir Samuel (Wanjarri) and Laverton-Leonora (Erlistoun) Study Areas: sampling dates and methods*. 1. Wanjarri Survey Area.

Fauna 14-21 Feb, 14-20 May, 24-29 Aug, Lithological Code** 1979 1980 1981 Surface

1W-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpv/Qqz 1W-02 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqc 1W-03 Q(M,B,R) Q (M,B,R) Q(M,B,R) Qps/Qqz 1W-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Czo+Qpv lW-OS Q(M,B,R) Q (M,B,R) Q(M,B,R) Qqz 1W-06 F,B,R F,M,B,R F,M,B,R Qps 1W-07 F,B,R F,B,R Qps 1W-08 F,B,R F,B,R Qps

2. Erlistoun Survey Area.

Fauna 22-28 Feb, 8-13 May, 18-23 Aug, Lithological Code** 1979 1980 1981 Surface

lE-01 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpk 1E-02 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qrm lE-03 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qrd 1E-04 Q(M,B,R) Q(M,B,R) Q(M,B,R) Qps/Qqz lE-OS Q(M,B,R) Q(M,B,R) Q(M,B,R) Qpv lE-06 F,B,R F,B,R Qps

*M = Mammal trapline (9 Elliotts, 9 Break-backs, 3 Cages) B = Bird searches R = Reptile searches F = Fenced pit line of six pits (4 shallow, 2 deep) with ca. 50 m drift fence Q = Fauna quadrat (fenced pit lines and, adjacent, a 200 metre x 200 metre bird quadrat) that was sampled daily for five days as described in Biological Surveys Committee of Western Australia (1984). ** The inventory prior to Appendix 1 lists the vegetation site codes and corresponding fauna sites.

51 20844, 20845, 20865-20867, 20886, 20904-20910, 23501-23610, 38654-38660 and 39968. Species nomenclature in this report follows that of the Western Australian Museum. The quadrats within the Erlistoun survey area in the Laverton-Leonora (LL) Study Area (Figures 3 and 5) allowed lithological units low in the landscape to be sampled, whereas the quadrats in the Wanjarri survey area (within Wanjarri Nature Reserve) in the Sandstone-Sir Samuel (SS) Study Area were positioned to sample units high in the landscape (Figures 2 and 4). Table 1 lists the surface lithology and associated "vegetation types" known from the two Study Areas, while Appendix 1 indicates those that were sampled for vertebrates, cross­ references the vertebrate and vegetation sample site codes, and describes the location, vegetation, floristics and substrate ofeach vertebrate site sampled. Only the main vegetation types of the most extensive landform units were surveyed for vertebrates (Broad Valleys, Sal.t Lake Features, Sandplains, Dunefields and Breakaways). This meant, for instance, that we sampled red sandplains (surface lithology = Qps) in the context of Broad Valleys, Dunefields as well as Sandplains (Appendix 1). It also meant that less extensive landforms such as Granitic, Banded Ironstone and Greenstone Hills were not surveyed at all, even though they are prominent in the Study Areas. We surveyed the vertebrate assemblage from at least one site in five of the 13 landform .units, eight of more than 70 surface lithologies, and in 12 of over 70 vegetation types known in the two Study Areas (Tables 1 and 2). Much of the floristic diversity of these surface-types was remote from our quadrats. Considering how poorly the vertebrate sampling addressed the environmental heterogeneity of the Study Areas, and how geographically localised our quadrats were (Figures 2 and 3), we encountered a surprisingly high proportion of its vertebrates (Table 3). This coverage was achieved for passerine birds (61 %) and small ground-dwelling mammals (81 %) because most species occurred on a variety of lithologies. For instance, the "average" passerine bird occurred in a mean of 3.7 (S.D. =2.3, n =44), and an "average" small ground-

Table 3 Proportion of the Sandstone-Sir Samuel and Laverton-Leonora Study Areas' known vertebrate species recorded on quadrats.

Number ofSpecies Total On quadrats' (%)

Birds passerines 72 44 (61) non-passerines 78" 19 (24) Reptiles snakes 16 1 (6) lizards 59c 39 (66) Amphibians 4C 3 (75) Smallb ground-mammals 11 9 (81)

* lWOl-05 (Wanjarri) & lEOl-05 (Erlistoun), excluding additional sample sites lW06-8 & lE06 (see Table 2). " Includes 30 water birds. b Indigenous species with an average adult body weight of less than 35 g (see Burbidge & McKenzie 1989). c Excluding species that only reach the periphery of the Study Areas (see reptile text).

52 (a) WANJARRI

10 9 9

8 ~ 7 U ~ 6 (f) LLo 0:: u.J ID ~ ~ 3

2

8 (f) 7 u.J U u.J a.. 6 (f) LL 5 0 5 0:: u.J ID 4 ~ :::l Z

2

22 23 24 25 26 27 8 10 11 12 18 19 20 21 22 DATE FEB 1979 MAY 1980 AUG 1981 No OF DAYS Figure 8 Species accumulation curves for small ground mammals recorded at sample sites in the a) Wanjarri (I WO I-I W06) and b) Erlistoun (I EO I-I E06) survey areas.

53 Table 4 Species of mammals recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areas indicating number of records at each vertebrate quadrat and sample site during each survey period. The three survey periods - Summer (February 1979), Autumn (May 19S0) and Winter (August 19SI) are indicated in columns I, 2 and 3 respectively for each quadratlsite'.

Vertebrate QuadratlSite lE' IEOI IE02 IE03 IE04 IEOS IE06 Vegetation Site' LLI LL2 LL3 LL4 LLS LL6 Stratigraphy Qpk Qnn Qrd Qps/Qqz Qpv Qps Drift Fence Nights SSSSSSSSS S ** S S S 0 2 S TACHYGLOSSIDAE Tachyglossus aculeatus X' X'O DASYURIDAE Antechinomys laniger Ningaui ridei 2. 0 .3. 010 Pseudantechinus wooleyae Sminthopsis crassicaudata o 0 S. hirtipes 020 S. macroura 2. 0 0 (2) I 0 S. ooldea 010 002 MACROPODIDAE Macropus robustus M. rufus 200 001 MOLOSSIDAE Mormopterus planiceps' Tadarida australis VESPERTILlONIDAE Chalinolobus gouldii Eptesicus finlaysoni Nyctophilus geoffroyi :1 0 0 Scotorepens balstoni MURIDAE Leporillus ?apicalis NO/omys alexis 001 Pseudomys hermannsburgensis 0 0 1 010 (4):1 2 S 2 1 o 0 001 Mus musculus 2.0142.2 111 200 I 1 CANIDAE Vulpes vulpes o 0 X' FELIDAE Felis ca/us 200 BOVIDAE 80S taurus Ovis aries CAMELlDAE Camelus dromedarius 00 X' o X'O EQUIDAE Equus caballus LEPORIDAE Oryctolagus cuniculus 300 x' 0 0

Sampling methods and dates for quadrats (IEOI-OS, IWOI-OS) and sample sites (IE06, IW06, IW07 and IWOS) are outlined in Table 2. See Appendix l. Opportunistic records (not on quadrats). Total observations during F79 (February 1979), MSO (May 19S0) and ASI (August 19SI). * These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (IE04 and IW04).

54 lW' IWOI IW02 IW03 IW04 IW05 IW06 IW07 IW08 SSI8 SS20 SS21 SS 1,2 S519 SS22 SS23 Qpv/Qqz Qqc Qps/Qqz Czo+Qpv Qqz Qps Qps Qps TOTALOBS.' 5 5 5 5 5 5 5 5 5 5 • 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 M80 A81

----~~~

X'O 0 X' X' 0

0 0 2 () fl () () I 2 0 () ..2 () () 2 0 7 9 0 0 1 2 () 0 1 0 0 ..2 0 0 1 0 0 1 0 0 2 3 2 0 1 12 4 7 0 I 0 0 1 0 7 4 1 0 Q 2 - 1 0 0 9 2 2 5- ..2 I 4 5 2 4 9 12 12 () II () 1 0 I 1 3 6

8 4 0 0 0 () 0 0 6 () 0 () () 8 10 8 la 6 0 0 3. () () () 0 () 0 29 22 25

Q) 0 0 4 1 0 0 6 3 0 7 4

(7) 311 0 0 I 0 0 9 3 16 0 1 2 () 0 1 1 2 ~) 0 H 9 0 13 (la) 1 2 () 0 19 1 8

X' X' 0 0 I 0 0 ..2 I 2 Q 2 0 - 0 1 1 4 10 1 0 0 0 2 3 I 5- 2 2 2 0 2 0 I 4(2) () 1 I 17 19 25 0 0 10 4 6

X' 0

2 0 0

0 ~ 0 0 ~ 0 10 0 0 o 30 () 10 30 0

0 X'O 0 1 0 0 X'

0 0 0 0

3 0 0

Faeces (Tachyglossus). Den (Vulpes). Warren in use (Oryctolagus). Tracks (Camelus). lS Sample includes sub-adults. (,) Sample includes post-partum, lactating or pregnant individuals. The form that ranges across southern Australia. Remains of old nest under breakaway.

55 (a) WANJARRI

40 37 35 3&

30 . ·3t· (j) UJ U UJ 25 Il. (j) LL.. 0 20 Cl:: UJ IJ) 15 ::2; ::::>z 10

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25 DATE FEB1979 I MAY 1980 I AUG1981 ISEP92 No OF DAYS 7 12 17 18

(b) ERLlSTOUN

40 38

35 35

30 (j) UJ U UJ 25 Il. (j) LL.. 0 20 Cl:: UJ IJ) 15 ::2; ::::>z 10

5

0+--+-+...... ,f-+--r--+-4-+-+-+-+---1-t--t--r---r-j...... ,...---1 22 23 24 25 26 27 28 7 8 9 10 11 12 13 18 19 20 21 22 28 DATE FEB 1979 I MAY 1980 I AUG 1981 I SEP 92 No OF DAYS 7 14 19 20 Figure 9 Species accumulation curves for reptiles (excluding amphibians) recorded at all sample sites (including opportunistic records) in the a) Wanjarri and b) Erlistoun survey areas.

56 mammal in 3.4 (2.2, 9), of the eight distinct Iithological surface-types sampled (Qpk, Qrm, Qrd, Qps, Qpv, Qqc, Czo and Qqz). In contrast, the "average" lizard was recorded on only 1.8 (1.1, 40) of these Iithological units; the 66% coverage for lizards may have been achieved because only certain lithologies (especially the widespread Qqz and Qps on which our sampling focussed) appear to have rich, compositionally distinct, lizard faunas (see Table 7). Species accumulation curves, in relation to days of effort, were used to investigate whether further sampling effort in the quadrats would have been cost-effective. Separate curves were compiled for reptiles and small ground mammals at the Wanjarri and Erlistoun survey areas. Although the curves for survey areas are clearly beyond a final detlection point (Figures 8 and 9), the reptiles of surfaces such as Qrd were clearly under-sampled (Figures 10 and 11). The seasonally tiered pattern general to the curves indicates the importance of sampling during several seasons. The results of the single day of reptile sampling in September 1992 imply that a longer session in spring would have added species. For example, if Spring records are removed from the Kurnalpi-Kalgoorlie quadrat lists (McKenzie et at. 1992), nearly half the species list of ground-dwelling vertebrates is eliminated. The analysis package PATN (Belbin 1989) was used to seek patterns of species composition in the data matrices. The clustering techniques selected were described in McKenzie et at. (1991 b). The species assemblages from the quadrats were used as the input data. The presence and absence of species on the quadrats, rather than their relative abundance, was used because limitations in sampling techniques, aggravated by staff and time limitations, precluded reliable estimates of relative abundance (Austin 1984, McKenzie et al. 1991a). Previous studies have shown that our broad-scale sampling techniques are also unsuitable for snakes, monitors and non-passerine birds, so they were excluded from the analyses (McKenzie 1984, McKenzie et al. 199Ia).

Table 5 Bat species recorded from the Wanjarri and Erlistoun survey areas during the survey of the Sandstone­ Sir Samuel and Laverton-Leonora Study Areas.

~ SITE': 2 3 4 5 6 I 8 Tadarida australis * ** Mormopterus planiceps * * + * Chalinolobus gouldii * ***** * Scotorepens balstoni *** *** * Eptesicus finlaysoni *** Nyctophilus geoffroyi * * * , I = 27"17'5 120"40'E (granite outcrop); 2 =27°20'5 120"40'E (Bullock Well, Mulga); 3 =27"32'5 120"43'E (Dingo Pool, Mulga); 4 =27"21'5 120"36'E (breakaway pool at 1W04); 5 27"24'5 120"41 'E (red sand dune near IW06); 6 27"24'5 1200 39'E (Homestead Dam, Mulga). 7 =28"16'5 122"06'E (lake Ikm north of 1E03). 8 =28°22'5 I 22"OTE (dam, Mulga). + =recorded on 25 September 1992 from sonar signature using ANABAT 11 (Corben 1991).

57 Qqc(1W02)

20 "I. 17

~ 14 U ~ 12

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25 DATE FEB 1979 MAY 1980 I AUG 1981 I SEP 92 No OF DAYS 12 17 18

CZQ & Qpv (1W04) 20 18,.

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25 DATE FEB 1979 I MAY 1980 I AUG 1981 ISEP 92 No OF DAYS 7 12 17 18

Qpv (1W01) 20

18 "

14 15 16 17 18 19 20 21 14 15 16 17 18 24 25 26 27 28 25 DATE FE81979 I MAY 1980 I AUG 1981 ISEP 92 No OF DAYS 7 12 17 18 58 Qps (1W03 & 1W06)

'0 20

18

U) UJ (3 g: 12 U) 10 0 10

1iS'" 8 ::;; ::J Z

14 15 16 17 16 19 20 21 14 i5 16 17 18 24 25 26 27 28 25 FEB 1919 MAY 1980 12 \ AUG 1981 171 s~: 92 7 !

Qqz (1W05)

20

18

" U) UJ Qqz (3 " I---1W05I UJ "- 12 1/) "- 10 10 10 0 10

UJ '"lJJ ::;; 1 ::J Z 4 5 4

14 15 16 17 18 19 20 21 14 i5 16 17 18 24 25 26 27 28 25 DATE FEB 1979 MAY 1960 AUG 1981 92 No OF DAYS 121 171 s;;

~ .. Figure 10 Species accumulation curves for reptiles (excluding amphibians) recorded on each surface sampled in the Wanjarri survey area. Accumulation curves for the individual sample sites on each surface are also included for comparison.

59 Qpv (1E05)

2

(f) w ,. uw a.. 12 1 Qpv f (f) . --lEOS u. 0 10 '"W • ::;"' 7 '"z 6 oL4=~~:re:::: 22232<4252627287 8 9 10 11 12 13 18 19 20 21 22 28 DATE FEB 1979 MAY 1980 I AUG 1981 I SEP 92 NoOF DAYS 14 19 20

Qrd (1E03) 20,. ,. 17 ~ I. U ~ Qrd 12 . I---1E03I (f) ~ 10 9 :,g'" • ::; '"Z 4

2324252627287 8 9 10 11 12 13 16 19 20 21 22 28 DATE FEB 1979 MAY 1980 I AUG 1981 I SEP 92 No OF DAYS 14 19 20

Qps (1 E04) 2.

22 20,. (f) w ,. U w a.. \4 (f) u. 0 12 11 '"w 10 ::;"' '"z

2324252627287 8 9 10 11 12 13 18 19 20 21 22 28 DATE FEB 1979 MAY 1980 I AUG 1981 I SEP 92 No OF DAYS 14 19 20

60 Qrm (1E02)

20

18

16

~ ~--1E02

8 9 10 11 12 13 18 19 20 21 22 28

DATE FEB 1979 MAY 1980 141 AUG 1981 19 i NoOF DAYS Si: 92

Qpk (1E01)

20

18

16

if) W 14 (3 w a. 12 I Qpk I if) --1E01 "- 0 10 9 w '"co 8 ::;; ::J Z

22 23 24 25 2f5 27 28 7 8 9 10 11 12 13 18 19 20 21 22 28 DATE FE81979 MAY 1980 I AUG 1981 ! SEP 92 No OF DAYS 71 14 19 I 20 A ~ Figure 11 Species accumulation curves for reptiles (excluding amphibians) recorded on each surface sampled in the Erlistoun survey area. Accumulation curves for the individual quadrats on each surface are also included for comparison.

61 Table 6 Data matrix of small ground-dwelling mammals re-ordered according to UPGMA classification outputs (Belbin 1989). The quadrats were classified according to similarities in their species composition, and the species according to their co-occurrence at the same quadrats and sample sites.

QuadratlSite Codes!

II 11 I I1I 1I 1I e e e e wwww ew ew 0000 0000 00 00 2135 1253 44 66 Species Pseudantechinus wooleyae * Sminthopsis hirtipes ** Notomys alexis **** Ningaui ridei **** ** ** Antechinomys laniger *** * Pseuomys hermannsburgensis ******* ** ** Sminthopsis ooldea ** ** ** Sminthopsis macroura * ***** Mus musculus * ** * ** Sminthopsis crassicaudata *** Lithological origin2 QQQQ QQQT TT TT

! Quadrat & sample site codes are printed vertically. Q = Quaternary; T = Tertiary.

Mammals Eighteen species of native mammal were extant on the quadrats during the survey (Table 4), These comprised two species of kangaroo, 9 small ground-dwelling mammal species (7 dasyurids and 2 rodents), the Echidna, and six species of insectivorous bat Eight species of introduced mammal were also encountered (including Mus musculus). In addition, more than 40 Goats (Capra hircus) were seen in October 1985 (CALM files), and several herds of 20 or more were seen by the authors in September 1992. According to Western Australian Museum records, 15 indigenous and two introduced mammals were previously known from the Study Areas. Two of the indigenous species were not recorded during our field work: Macrotis lagotis (M632, Laverton, 1924) and Myrmecobius fasciatus (M351, Ida Hill Mine, 1918, 28"41 'S 122"24'E); both are known to have become extinct or very much rarer throughout most of their former range since European settlement (Burbidge and McKenzie 1989). Another 14 native species have been identified from bone material in exposed, superficial sub-fossil deposits in the Youanrni-Leonora Study Area (Figure 1), immediately to the south of the Study Areas described herein (Appendix 13 in Henry-Hall 1990). Twelve of these species are known to have become extinct or very much rarer throughout mainland Australia since European settlement (Burbidge and McKenzie 1989): Dasycercus cristicauda,

62 10

9 8 6 5 4

8 3 7 7 4

5 9 10 2 6 2 (a) 3 (b) 11

Figure 12 Three-dimensional ordinations of the small, ground-dwelling mammal data from sites at Wanjarri (lWOI-lW06) and Erlistoun (lEOI-IE06). Minimum Spanning Trees have been superimposed. Shaded wedges represent projections into the 3rd dimension. (a) Species ordinated according to their co-occurrences at the same sites. Stress=0.037. (b) Sites ordinated according to similarities in their species composition. Stress=0.07. See text for details of analyses. a) Species ordination. b) Site ordination. KEY: KEY: I. Sminthopsis crassicaudata Red Tertiary sand. hi~h in the landscape: 2. Mus musculus 1. lE06 & 1W06: as dunes (Qps) at the top of the landscape. 3. Sminthopsis macroura 2. lW03: as sheets mantling Quaternary soil plains (QpslQqz). 4. Sminthopsis ooldea Quaternary surfaces at middle levels in the landscape: 5. Pseudomys hermannsburgensis 3. lWOS: Soil plain (Qqz). 6. Antechinomys laniger 4. lW02: Gibber slope (Qqc). 7. Ningaui ridei 5. lW01: Alluvium (Qpv) in freshwater drainage line incising the Qqz. 8. Notomys alexis 9. Sminthopsis hirtipes Quaternary surfaces lower in the landscape 10. Pseudantechinus wooleyae 6. lEOS: Alluvium (Qpv) in freshwater drainage line incising a QpslQqz surface. 7. lE03: Brownish sand (Qrd) peripheral to Qnn. 8. lE02: Qccluded drainage of saline alluvium (Qnn) at the bottom of the landscape. 9. lEOl: Gypsum dune (Qpk) peripheral to Qnn. Tertiary surfaces. medium to low in the landscape 10. lE04: Red sand sheet, mantling Quaternary soil plains (QpslQqz). 11. 1W04: As duricrust breakaway complex (Czo) and associated Quaternary alluvial deposits (Qpv).

63 Table 7 Species of reptiles recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areas indicating number of records at each vertebrate quadrat and sample site during each survey period. The three survey periods - Summer (February 1979), Autumn (May 1980) and Winter (August 1981) - are indicated in columns 1,2 and 3 respectively for each quadratlsite'.

Vertebrate QuadratlSite lE' IEOI IE02 IE03 IE04 IEOS IE06 Vegetation Site2 LLl LL2 LL3 LL4 LLS LL6 Stratigraphy Qpk Qrm Qrd Qps/Qqz Qpv Qps Drift Fence Nights 555 555 5 5 5 5 ** 5 5 5 o 2 5 Cye/orana maini 040 0 0 Cye/orana platycephala 200 Neobatrachus ?kunapalari 150 0 0 2 0 0

Diplodactylus conspicillatus 100 0 0 D. elderi S D. pulcher D. squarrosus 100 100 D. strophurus 100 D. wellingtonae 010 Gehyra purpurascens 001 S G. variegata o 223 S S 0 0 Heteronotia binoei 113 Underwoodisaurus milii 100 Rhynchoedura ornata 101

Delma butleri o o D. nasuta S o o 3 0 Lialis burtonis 100

Caimanops amphiboluroides Ctenophorus caudicinctus C. fordi 2 0 3 C. inermis 1 1 0 0 I 0 C. isolepis 2 3 2 2 2 0 - I 0 C. reticulatus 2 0 0 C. salinarum 1 0 2 0 0 I 2 I 2 C. scutulatus 1 0 0 0 0 Moloch horridus 1 0 2 Pogona minor 0 0 2 0 0 I 0 I S

, Sampling methods and dates for quadrats (1E01-05, IWOI-05) and sample sites (lE06, IW06, IWO? and IW08) are outlined in Table 2. 2 See Appendix I. ) Opportunistic records (not on quadrats). 4 Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981); excludes records from October 1987 and September 1992 visits. * These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (IE04 and IW04). S Recorded in September 1992.

64 lW' lWOl lW02 lW03 lW04 lW05 IW06 lW07 lWOS SSlS SS20 SS2l SSl,2 SSl9 SS22 SS23 Qpv/Qqz Qqc Qps/Qqz Czo+Qpv Qqz Qps Qps Qps TOTALOBS" 5 5 5 5 5 5 5 5 5 5 * 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 MSO ASl I 0 0 2 0 0 I 0 0 5 4 0 I 0 0 0 0 4 0 0 5 0 0 7 0 0 4 0 0 0 0 0 I 0 2 0 20 9 0

0 0 0 0 3 0 I 7 0 S 0 0 2 0 0 3 0 0 2 0 0 - 0 1 I 0 I 0 0 1 0 0 0 0 0 0 4 I 0 0 0 I I 0 S 7 0 1 8 3 25 0 I 2 0 0 4 I 4 I 0 0 0 0 4 0 0 6 0 1

1 0 0 I 1 0 0 0 1 - 0 1 0 4 2 I 0 0

I 0 0 0 1 0 II 0 0 0 2 1 0 2 I 0 4 2 0 3 6 0 0 1 0 0 I 0 0 0 0 10 2 0 2 0 0 3 0 0 2 0 0 0 1 II 6 3 2 0 0 0 5 0 I I 0 0 4 I 5 2 0 0 I 0 0 2 0 2 2 0 0 3 0 3

65 Table 7 (conI.)

Vertebrate QuadratfSite lE' IEOI IE02 IE03 1E04 IE05 IE06 Vegetation Site2 LLl LL2 LL3 LL4 LL5 LL6 Stratigraphy Qpk Qrm Qrd Qps/Qqz Qpv Qps Drift Fence Nights 555 555 555 5 ** 555 025

CryplOblepharus carnabyi 120 C plagiocephalus S Ctenotus ariadnae C calurus C grandis C greeri 200 C helenae 120 120 C leonhardii C pantherinus S 3 2 C quattuordecimlineatus C schomburgkii o 2 Egernia depressa E.jormosa Eremiascincus richardsonii Lerista bipes L. deserlOrum 001 S S S L. muelleri 004 o 0 100 Menetia greyii 002 S S Morethia butleri 002 Tiliqua multijasciata 100 T. occipitalis o 1

Varanus brevicauda 100 100 V. caudolineatus 010 001 V. gouldii 100 V. panoptes

Furina ornata 1 0 Pseudechis australis o 0 Pseudonaja nuchalis Ramphotyphlops hamatus 100 Vermicella bertholdi s

Species Richness 33 7 3 9 12 8 2

I Sampling methods and dates for quadrats (1E01-05, IWO 1-05) and sample sites (I E06, IW06, IW07 and IW08) are outlined in Table 2. 2 See Appendix I. , Opportunistic records (not on quadrats). 4 Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981); excludes records from October 1987 and September 1992 visits. * These should read 10 because an additional fenced pit line was set adjacent to the bird quadrat (lE04 and IW04). S Recorded in September 1992.

66 lW' IWOI IW02 IW03 IW04 IW05 IW06 IW07 IWOS 551S 5520 5521 551,2 5519 5522 5523 Qpv/Qqz Qqc Qps/Qqz Czo+QpvQqz Qps Qps Qps TOTALOB5' 5 5 5 5 5 5 5 5 5 5 * 5 5 5 5 4 5 5 0 2 5 0 3 5 F79 MSO ASI

2 0 S 3 0 3 I 0 0 0 0 I 0 0 0 I 0 0 2 0 0 0 0 I 2 0 0 4 4 I 2 0 0 5 0 0 I 0 0 2 0 0 10 0 0 0 I I 0 3 3 2 I 0 0 0 2 I 0 2 I 0 0 5 0 I 0 0 2 0 0 2 I 0 11 3 2 2 0 0 0 0 3 0 0 2 0 0 2 0 0 I 0 0 1 0 0 5 I 0 0 1 0 0 0 0 1 5 5 1 0 5 I 1 I 0 0 3 0 0 5 1 3 0 I 0 I 3 I 0 0 2 0 0 I 0 I

1 0 0 2 0 0 I 0 2 3 I 3 I 0 0 I 0 0 I 0 0

1 I 0 0 1 0 1 0 0 1 0 0 0 0 2 0 0 S

23 10 7 II 14 6 6 2 5 57

67 Phascogale caIura, Chaeropus ecaudatus, Isoodon auratus, Trichosurus vulpecula, Onychogalea lunata, Leporillus apicalis, L. conditor, Notomys amplus, N. longicaudatus, Pseudomys fieldi and Rattus tunneyi. Except for the Dasycercus, there is no evidence that they persist in the district. CALM Departmental files include a letter from J.L. Bannister, dated 12 December 1969, noting that T.K. Moriarty had sent a female Dasycercus cristicaudata to the W.A. Naturalists Clubs Wildlife Show in 1967, but the specimen was subsequently lost. At that time, Bannister was the Curator of Mammals at the Western Australian Museum and Moriarty owned and operated the Kathleen Valley pastoral lease (= Wanjarri). G. Connell (pers. comm.) trapped D. cristicaudata in the south-western part of the Wanjarri Nature Reserve in 1991. Bones of Pseudomys holami and P. desertor were also found in the sub-fossil deposits from the Youanmi-Leonora Study Area. However, P. holami was not trapped during our survey, or in other Study Areas in the northern Goldfields even though appropriate habitats were sampled (Dell and How 1992, How and Dell 1992). Noting that the species was readily trapped in survey areas to the south (e.g. Dell and How 1988), the sub-fossil record from the Youanmi-Leonora Study Area may represent the northern limit of this species' range. P. desertor may still be extant in the Study Areas.

60 ~•..

50 I- .1979 01980 Cl) .1981 UJ 40 I- r' () UJ a. Cl) L.l. 0 30 I- f· f-. 0:: UJco ::lE 20 ---- ...... :::>z

10 f-· ..-

0 .d 2 3 4 5 6 7 8 9 10 11 12 13 14 15 NUMBER OF DAYS

Figure 13 Species accumulation histogram for reptiles (excluding amphibians), showing seasonal differences in the number of species encountered per day of sampling effort. Results from the Wanjarri and Erlistoun survey areas are combined.

68 The extant native mammal fauna comprises only Eyrean species. There arc several species in common with the adjacent deserts (the Helms and Keartland Botanical Districts): e.g. Sminthopsis ooldea. S. hirtipes. Ningaui ridei. Notomys alexis and Pseudomys hermannsburgensis. This retlects the mozaic of red sandplain and sand dune surfaces scattered through the northern Goldfields. The presence of Pseudantechillus wooleyae and Eptesicus jinlaysolli, rather than their desert counterparts (Pseudantechinus macdonnellellsis and Eptesicus baverstocki occur in the Helms District), is consistent with the phytogeographic position of our Study Areas in the Austin Botanical District, as discussed in Keighery et al. (this publication). Bats were sampled using mist nets, a bat trap and spotlight shooting. Sites were usually on

Table 8 Amphibians and reptiles from the Sandstone-Sir Samuel Study Area not recorded during the survey. ~------_._-_. ------_.._._--_.__._--- Species Specimen* Source Location

Limnodynastes spenceri WAM R 104473 Yakabindie Pseudophryne occidenlalis WAM R 100534 Barr Smith Range Diplodactylus elderi WAM R 26878 Montague Range D. squarrosus WAM R 104471 Yakabindie Nephrurus verrebralis WAM R 48031 Wanjarri N. wheeleri WAM R 8942 Sandstone Oedura marmorata WAM R 84357 Anketell Outstation Pygopus nigriceps WAM R 1103 Booylgoo Spring Ctenophorus ornatus WAM R48026 Wanjarri C. scutulatus WAM R 27219 Wanjarri Tvmpanocryptus cephala WAM R 100967 Wanjarri Cryptob!epharus plagiocephalus WAM R 82561 Yeelirrie Homestead Egernia inornala WAM R 48037 Wanjarri E. kintorei WAM R 27228 Wanjarri E. striata WAM R 100964 Wanjarri Lerisla deserrorum WAM R 19766 Wanjarri Tiliqua occipitalis WAM R 37786 Wanjarri Varanus eremius WAM R 27230 Wanjarri V. giganteus WAM R 27232 Wanjarri V. gouldii WAM R 12406 Wanjarri Morelia stimsoni WAMR 1417 Booylgoo Spring Acanrhopis pyrrhus WAM R 30599 Albion Downs Denisonia fasciata WAM R 40522 Wanjarri Pseudechis australis WAM R 12599 Wanjarri P. burleri WAM R 7627 Booylgoo Spring Pseudonaja modesta WAM R 1122 Booylgoo Spring Rhinoplocephalus monachus WAM R 19777 Wanjarri Vermicella bertholdi WAM R 31671 Wanjarri V. semirasciala WAM R 40523 Wanjarri Ramphotyphlops waitti WAMR 19778 Wanjarri

*WAM Western Australian Museum, Perth

69 pools, places where bats congregate to drink. The observed assemblages are listed in Table 5. A solitary N. geoffroyi was found roosting in a Mulga spout during the day. Tadarida australis seldom foraged less than 10 metres above the ground, and always well clear of the vegetation canopy. It was often recorded in pairs. Mormopterus planiceps and Chalinolobus gouldii foraged just above the open woodland canopy (6 9 metres above the ground), although the C. gouldii flew more slowly and often foraged between tree canopies. Scotorepens balstoni and Eptesicus jinlaysoni were seen foraging the gaps between trees and shrubs, although E. jinlaysoni flew more slowly and closer to the vegetation, with frequent changes in direction. Two Nyctophilus geoffroyi were tracked at night using luminescent Cyalume beads (see Buchler 1976) in a low open Mulga (Acacia aneura) woodland near the Wanjarri homestead. They fluttered slowly along less than two metres above the ground, often changing direction and occasionally diving almost vertically to its surface. These differences in bat foraging micro-habitats are consistent with wing-shape differences described for these species by Fullard et al. (1991) and McKenzie and Start (1989). Data on small ground mammals collected during the trapping program are summarised in Table 4. The trap-effort values at the top of this table provide a relative measure of sampling effort because all small ground mammals recorded on quadrats were trapped. The results of the numerical classifications are presented as a re-ordered data matrix in Table 6. To confirm the robustness of the clusters, species and quadrats were ordinated in 3­ dimensional space using Semi-strong Hybrid Scaling (Belbin 1991), then a Minimum Spanning Tree was superimposed (Figure 12). These analyses revealed a gradient in the composition of the small ground mammal assemblages in the survey areas that could be explained in terms of surface lithology. The upper levels in the landscape are Tertiary surfaces. These were characterised by Parantechinus wooleyae where duricrust is exposed as breakaways along eroding edges of the uplands, and by Smimhopsis hirtipes on the red sand dune systems (Figure 12 and Table 6). The Tertiary surfaces also supported Notomys alexis, Ningaui ridei, Sminthopsis ooldea and Pseudomys hermannsburgensis, but these four species also occur down the landscape sequence to levels where sheets of Tertiary sands have mantled, or are mixed into the upper profile, of heavier Quaternary surfaces. The last two of these "Tertiary" species extend into Quaternary sequences, although Sminthopsis ooldea avoids the heavier soils. The lower levels in the landscape are Quaternary surfaces. Antechinomys laniger favoured the heavy soils of Quaternary plains, away from the salt lake systems. Sminthopsis macroura was found on all Quaternary surfaces except the salt lake bottoms and gypsum dunes. At the bottom of the landscape, the salt lake surfaces and their peripheral features supported the S. crassicaudata and Mus musculus populations. A similar pattern in composition was noted in the Kurnalpi-Kalgoorlie Study Area, although results from Kurnalpi indicate that S. crassicaudata can disperse across Tertiary surfaces (McKenzie et al. 1992). Although a session of sampling in spring would have captured additional species on quadrats, the levels of sampling were sufficient to show that compositional gradients reflected the geomorphological sequences in the landscape's surface stratigraphy (see Figure 12).

Reptiles and Amphibians Three frogs and 54 reptile species were recorded in the Sandstone-Sir Samuel and

70 Table 9 Amphibians and reptiles from the Laverton-Leonora Area not recorded during the survey.

Species Specimen' Source Location

Liml1odYl1astes spel1ceri WAM R 1248 Laverton Pseudophryl1e occidel1talis WAM R 21536 White Cliffs Diplodactylus assimilis WAM R 20651 White Cliffs D. gral1ariel1sis WAM R 97289 60km SE Leinster D. pulcher WAM R 1246 Laverton D. stel10daclylus WAM R 85633 Pt Salvation Nephrurus levis WAM R 85634 39km E Laverton N vertebralis WAM R 78156 10km S Leonora Pygopus l1igriceps WAM R 85592 39km E Laverton Ctel1ophorus caudicil1ctus WAM R 25946 40km NE Laverton Tvmpal10cryptus cephala WAM R 1240 Laverton Ctel10tus ariadl1ae WAM R 85714 39km E Laverton C. calurus WAM R 85781 39km E Laverton C gral1dis WAM R 86217 39km E Laverton C hal1lol1i WAM R 86453 Laverton C leol1hardii WAM R 1234 Laverton C pial1kai WAM R 85690 39km E Laverton C qualluordecimlil1eatus WAM R 89073 39km E Laverton C severus WAM R 20617 Ilkm E Leonora Egernia depressa WAM R 1200 Laverton Eformosa WAM R 20659 White Cliffs E il10rnata WAM R 86081 39km E Laverton E richardi WAM R 25945 40km NE Laverton E striata WAM R 86136 39km E Laverton Eremiascil1cus richardsol1ii WAM R 17897 Cosmo Newbery Lerista bipes WAM R 85884 39km E Laverton Varal1us eremius WAM R 11609 Laverton V pal10ptes WAM R 98075 Bundarra Station V tristis WAM R 100396 39km E Laverton Morelia stimsol1i WAM R 13852 Cosmo Newbery Demal1sia psamophis WAM R 87884 Cosmo Newbery Del1isol1ia fasciata WAM R 1426 Laverton Pseudechis bUileri WAM R 22395 Laverton Pseudol1aja modesta WAM R 15052 Cosmo Newbery P. l1uchalis WAMR11047 Leonora Rhil1oplocephalus mOl1achus WAM R 11475 Laverton Vermicella bertholdi WAM R 85353 White Cliffs Station V fasciolata WAM R 1751 Laverton V semi/asciata WAMR 13854 Cosmo Newberv Ramphotyphlops el1doterus WAM R 13855 Cosmo Newbery R. waitii WAM R 21336 Laverton

,WAM Western Australian Museum, Perth

7\ 10

3

5a 5b

Figure 14 Three-dimensional ordination of the reptile data (except snakes and moniters) from sites at Wanjarri (lWOI-IW08) and Erlistoun (lEOI-IE06). Stress=0.14. Sites are ordinated according to similarities in their species composition. A Minimum Spanning Tree has been superimposed. Shaded wedges represent projections into the 3rd dimension. See text for details of analyses. KEY: Ouaternary soils low in the landscape I. lE02: Occluded drainage of saline alluvium (Qrm). 2. lE01: Gypsum dune (Qpk) peripheral to Qrm. 3. lE05: Alluvium in freshwater drainage line (Qpv) incising a QpslQqz surface. 4. lE03: Brownish sand (Qrd) peripheral to Qrm. Red Tertiary sand Sa-d. lW06, lWOS, lE06 & lW07 (respectively): as dunes at the top of the landscape (Qps). 6a-b. lE04 & lW03 (respectively): as sheets mantling Quaternary soil plains (QpslQqz). Ouaternary soils at middle levels in the landscave 7. lW05: Soil plain (Qqz). 8. lW02: Gibber slope (Qqc). 9. lW01: Alluvium (Qpv) in freshwater drainage line incising soil plain (Qqz). Tertiary breakaway complex 10. lW04: Duricrust outcrop (Czo) and associated Quaternary alluvial deposits (Qpv).

72 Laverton-Leonora Study Areas during our survey. The reptile species comprised 11 geckos, 3 legless lizards, 10 dragons, 21 skinks, 4 goannas and 5 snakes. These are listed in Table 7. A computer search of the W.A. Museum collection revealed records of another 2 amphibian and 31 reptilian species from the Study Areas during the period of European settlement (Tables 8 and 9). Thus, a minimum of 5 amphibian and 85 reptile species were extant in these Study Areas at the time of European settlement: 78 reptiles in the Laverton-L,e(mora and 64 in the Sandstone-Sir Samuel Study Area. However, one of the amphibians and 10 of the lizards listed in Tables 8 and 9 are only known from the periphery of our Study Areas; their geographic ranges are centred elsewhere (Pseudophryne occidentalis, Diplodactvlus assimilis, D. granariensis, D. stenodactvlus, Nephrurus ICl'is, N. Ivheeleri, Oedura marmorata, Ctenotus piankai, C. severtls, Egernia kintorei and E'. richardi). The proportion of the known reptile species richness that we recorded from quadrats is listed in Table 10. Clearly, our survey methods were less effective for monitors and snakes. Similar biases in reptile sampling have been discussed by McKenzie et al. (1987). The rich reptile lists from the Sandstone··Sir Samuel and the Laverton-Leonora Study Areas reflects long-term collecting by Eric Pianka (Pianka 1969, 1989) at sites in the eastern portion of our Study Areas, and by T.K. Moriarty at Wanjarri. Their richness compares with the 66 species known from the Kurnalpi-Kalgoorlie Study Area to the south (McKenzie et al. 1992). Although they have not actually been recorded within the Study Areas, at least another 2 amphibian and 5 lizard species might be expected on the basis of their known distributions in Western Australia: Cyclorana australis, Neobatrachus wilsmorei, Gemmatophora longirostris, Ctenows uber, C. brooksi, C. colletti and C. dux. Aside from wide-ranging species, there are three more specialised components in the reptile fauna: (I) Species with distributions centred on the heavier soils of the arid pastoral lands to the north, south and west (e.g. Diplodactylus squarrosus, D. wellingtonae, Caimanops amphiboluroides, Ctenophortls salinarum, C. scutulatus, Ctenotus severus, Egernia depressa, E. formosa and Tiliqua occipitalis).

Table 10 Proportion of the Sandstone-Sir Samuel and Laverton-Leonora Study Areas' known reptile species richness recorded on quadrats, by family.

Known l From Quadrats2

12 geckoes 10 (83%) 4 legless lizards 2 (50%) 12 dragons 9 (75%) 24 skinks 17 (71%) 7 monitors 3 (43%) 16 snakes 1(6%)

75 TOTAL 42 (56%)

From both Study Areas, excluding peripheral species (see Tables 8 & 9). Excludes the additional sample sites IW06-8 & IE06 (see Tables 2 & 7)

73 (2) Species that favour the red sandplains and dunefields that dominate the Great Victoria Desert to the east, and are scattered across the two Study Areas (e.g. Ctenophorus isolepis, C. fordi, Ctenotus calurus, C. quattuordecimlineatus, Lerista bipes and L. desertorum). (3) A Bassian component with distributions centred on semi-arid districts, that comprises the amphibian Pseudophryne occidentalis and the lizards Diplodactylus granariensis, Phyllurus milii, Ctenophorus omatus and Egemia richardi. None of these extend into the wettest areas of mesic South-western Australia. The dominance of arid zone reptiles is consistent with phytogeographic patterns; the Study Areas are entirely within the Eremaean Phytogeographic Province. They are in the north­ eastern (inland) corner of the Austin Botanical District, and remote from the South-Western Interzone (Beard 1980). A more prolonged session of sampling in the springtime would have captured additional lizard species on quadrats; even a single day of sampling in the spring of 1992 revealed additional species at six quadrats (Table 7, Figures 9-11). Quadrats on equivalent surfaces in adjacent Study Areas were richer in species (Dell and How 1992, How and Dell 1992). Nevertheless, from Figure 13 it is clear that the summer sampling session was more than twice as effective as the winter and autumn visits. The richest quadrat for reptiles was 1W04, an ecotone between Czo (Tertiary laterite) and Qpv (a gully incising the plateau edge - see Appendix 1). The lizard assemblages at two of the quadrats on alluvial surfaces OWOS =Qqz, lW02 =Qpc) were clearly incomplete (Figure 10). Their richness may have been reduced by over-stocking; the Mulga tree canopy on these quadrats was dead and the surface of the soil was bare and sheet-eroded (see Appendix 1). Figure 14 presents a 3-dimensional scatterplot derived by ordinating the quadrats according to similarities in their lizard species composition, then superimposing a Minimum Spanning Tree (the analysis pathway described earlier in this paper). These numerical analyses reveal a strong relationship between the composition of lizard assemblages and surface lithology. The levels of sampling were sufficient to show that gradients in composition reflected the geomorphic sequences in surface stratigraphy across the landscape. Analysis of data from another Eastern Goldfields Study Area yielded a similar conclusion (McKenzie 1984). In contrast, rainfall and temperature attributes are usually found to be significant as scalars of lizard composition in study areas with significant climatic gradients, such as areas with greater relief or those that cover entire districts (e.g. McKenzie et al. 1987, How et al. 1987, Gambold and Woinarski in press).

Birds Ninety-three species of bird were recorded in the Sandstone-Sir Samuel and Laverton­ Leonora Study Areas during our survey. Sixty-three of these (Table 11) were recorded on surface-types consistently sampled by both quadrat and opportunistic techniques. Species known from the Sandstone-Sir Samuel and Laverton-Leonora Study Areas, but not recorded on quadrats during our surveys, are listed in Table 12. Additional birds have been recorded on adjacent salt lakes to the north of the Sandstone-Sir Samuel Study Area (Curry 1979, Storr 1985) and to the east of the Laverton-Leonora Study Area (Storr 1986). Species recorded on Lake Violet (26°39'S 120° 16'E) were: Blue-billed Duck, Eastern Golden Plover, Greenshank, Sanderling, Red-necked Stint, Long-toed Stint,

74 Pectoral Sandpiper, Curlew Sandpiper, Oriental Pratincole and Silver Gull. Three other birds recorded in adjacent areas to the north and cast of both Study Areas were the Samphire Thornbill, Calamanthus and Painted Finch (Storr 1985, 1986). Thirty-two of the 61 non-passerines listed in Table 12 arc swamp or lake birds; no quadrats were sampled in these habitats during our field work. Many of the other non-passerines in this table arc species near the edge of their geographic range (e.g. Regent Parrot, Purple-crowned Lorikeet and Elegant Parrot); others have declined since settlement (Malleefowl, Princess Parrot), or arc vagrants or seasonal migrants (e.g. Black-cared Cuckoo, Rainbow Bee-cater). Of the 27 passerine species that wc failed to record on quadrats, the Grey-crowned Babbler and Purple-gaped Honeyeater (further records arc needed to confirm this record) arc species at the limits of their range, species such as the Black Honeyeater, Grey Honeyeater and White­ browed Woodswallow arc vagrants or seasonal nomads, and the Western Wedgebill is now scarce through much of its former range. The White-plumed Honeyeater, the Orange Chat and the White-fronted Chat, Rufous-crowned Emu-Wren, Striated Grasswren and Spinifex­ bird favour habitats that wc did not sample using bird quadrats. The 151 species recorded since European settlement in the Sandstone-Sir Samuel and Laverton-Leonora Study Areas comprises 80 non-passerines and 71 passerines. The Eyrean component is dominant because of the geographical location of the Study Areas and their variety of distinctly Eremaean vegetations.

GEOLOGY* QUADRAT* SIMILARITY COEFFICIENT (CZEKANOWSKI) 0.16 0.29 0.42 0.55 0.68 0.81

Opk 1E01 Onn 1E02 I SALINE Ord 1E03 Ope 1W02 I Opv 1E05 Czo+Opv 1W04 I Opv/Oqz 1W01 Oqz 1W05 I ALLUVIAL Ops/Opz 1E04

Ops/Oqz 1W03 I RED SAND

* SEE TABLE 2

Figure 15 Bird quadrats classified according to similarities in their passerine species composition (excluding non-passerines and corvids) using the Czekanowski (1932) similarity algorithm and unweighted pair-group mean averaging (U.P.G.M.A.) as a sorting strategy (Belbin 1987).

75 Table 11 Species of birds recorded in the Laverton-Leonora (E) and Sandstone-Sir Samuel (W) Study Areas indicating number of sightings at each vertebrate quadrat during each survey period. The three survey periods - February 1979 (Summer), May 1980 (Autumn) and August 1981 (Winter) are indicated in columns I, 2 and 3 respectively for each quadrati.

Stratigraphy Qpk Qrm Qrd Qps/Qqz Qpv Vegetation Site2 LLl LL2 LL3 LL4 LL5 Vertebrate Quadrat IEOI lE02 lE03 IE04 IE05

NON-PASSERINES Emu T 2 - Hoary-headed Grebe Little Pied Cormorant White-faced Heron Pacific Heron Mountain Duck Grey Teal Black Duck Pink-eared Duck Wood Duck Collared Sparrowhawk Little Eagle Wedge-tailed Eagle 8 - - .J. - 1 - Spotted Harrier Brown Falcon - 1 Australian Kestrel - I - - .2 1 Australian Hobby Black-tailed Native-hen Eurasian Coot Australian Bustard Banded Plover Black-fronted Plover Red-capped Plover Common Sandpiper Black-winged Stilt Bush Stone-curlew Diamond Dove 8 Common Bronzewing 4 - Crested Pigeon - 2 5 4 Mulga Parrot 4 5 Each quadrat was sampled for five days over three seasons. See Appendix I. Opportunistic sightings (not on quadrats). Total observations during F79 (February 1979), M80 (May 1980) and A8I (August 1981). T Tracks x No season assigned for opportunistic sightings. .2 All sightings of birds in flight. * Recorded in August 1982. 99 Unspecified number of individuals (99+).

76 77 Table 11 (cont)

Stratigraphy Qpk Qrm Qrd Qps/Qqz Qpv Vegetation Site2 LLl LL2 LL3 LL4 LL5 Vertebrate Quadrat IEOI IE02 IE03 IE04 IE05 Ring-necked Parrot 2 - 3 1 2 2 Bourke's Parrot 3 2 I Budgerigar 2. ~ - ~ 1 I Cockatiel lQlQ 12. I 2. Galah 1 6 2 1 :l 20 :l Pallid Cuckoo Horsfield's Bronze-Cuckoo Tawny Frogmouth Australian Owlet-nightjar Red-backed Kingfisher

PASSERINES White-backed Swallow Tree Martin Richard's Pipit 3 - 3 2 Ground Cuckoo-shrike .1 ­ Black-faced Cuckoo-shrike 2 2 I White-winged Triller lQ - I Jacky Winter - 1 Hooded Robin - 2 1 Red-capped Robin - 5 7 Rufous Whistler - 1 Grey Shrike-thrush Crested Bellbird 6 1 10 - 2 Willie Wagtail - 2 - 3 - 1 Cinnamon Quail-thrush White-browed Babbler Southern Whiteface Weebill - 4 4 Broad-tailed Thornbill Yellow-rumped Thornbill Slaty-backed Thornbill Chestnut-rumped Thornbill - - 3 - 3 Striated Grasswren

Each quadrat was sampled for five days over three seasons. See Appendix I. Opportunistic sightings (not on quadrats). Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981). T Tracks x No season assigned for opportunistic sightings. 1 All sightings of birds in flight. * Recorded in August 1982. 99 Unspecified number of individuals (99+).

78 79 Table 11 (cont.)

Stratigraphy Qpk Qnn Qrd Qps/Qqz Qpv Vegetation Site2 LLI LL2 LL3 LL4 LL5 Vertebrate Quadrat IEOI IE02 IE03 IE04 IE05

Variegated Fairy-wren 2 White-winged Fairy-wren 2024 13 3425 10 Brown Songlark Rufous Songlark Australian Sittella White-browed Tree-creeper Mistletoebird 2 5 2 Striated Pardalote 2 Brown Honeyeater - I Black Honeyeater Pied Honeyeater 2 4 White-plumed Honeyeater Singing Honeyeater 10 4 6 Yellow-fronted Honeyeater 6 3 3 White-fronted Honeyeater - - 81 - -2.2 - - 40 - 231 2 15 Yellow-throated Miner -- 7 - 15 83927 2 - 22 Spiny-cheeked Honeyeater 210 - 4 I 3 17 12 Crimson Chat - 20 43 11 Zebra Finch - 22 - Magpie-lark Black-faced Woodswallow 338 2 -- I 6 - 17 1 Little Woodswallow Masked Woodswallow White-browed Woodswallow Pied Butcherbird 2 3 - 1 311 3 Grey Butcherbird Australian Magpie Grey Currawong Spotted Bowerbird Little Crow -lQ - Corvus spp - I 1: - 2 TOTAL SPECIES/SEASON 5 9 12 3 4 8 7 4 13 10 10 10 262421 TOTAL SPECIES/QUADRAT 17 12 22 19 40

Each quadrat was sampled for five days over three seasons. See Appendix l. Opportunistic sightings (not on quadrats). Total observations during F79 (February 1979), M80 (May 1980) and A81 (August 1981). T Tracks x No season assigned for opportunistic sightings. 2 All sightings of birds in flight. * Recorded in August 1982. 99 Unspecified number of individuals (99+).

80 81 Table 12 Bird species known from the Laverton-Leonora and Sandstone-Sir Samuel Study Areas that we did not encounter on our quadrats at Erlistoun (E) and Wanjarri (W). Sources include our own opportunistic records from Table II (this publication) (0), the RAOU Atlas of Australian Birds (Blakers et al. 1984) (a), Anon. (1993) (b), Moriarty (1972) (m), Pianka & Pianka (1970) (p) and Storr (1985, 1986) (s). (*) denotes unconfirmed records.

NON-PASSERINES Red-necked Avocet m B1ack-throated Grebe m Bush Stone-Curlew 0 Hoary-headed Grebe 0 Gull-billed Tern b Australian Pelican a Whiskered Tern Little Black Cormorant s Purple-crowned Lorikeet Little Pied Cormorant 0 Regent Parrot m White-faced Heron 0 Princess Parrot m Pacific Heron 0 * Elegant Parrot m Great Egret a Scarlet-chested Parrot m Straw-necked Ibis Major MitchelliEs Cockatoo s Black Swan m Black-eared Cuckoo m Mountain Duck 0 BoobookOwl m Grey Teal 0 Barn Owl m Black Duck 0 Tawny Frogmouth 0 Pink-eared Duck 0 Spotted Nightjar m Wood Duck 0 Rainbow Bee-eater m Musk Duck m Black-shouldered Kite m Black Kite m PASSERINES Square-tailed Kite m Welcome Swallow a Black-breasted Kite m Tree Martin 0 Whistling Kite a Fairy Martin m Brown Goshawk m White-backed Swallow 0 Little Eagle 0 Western Wedgebill a Spotted Harrier 0 Grey Fantail a * Marsh Harrier Grey-crowned Babbler m Peregrine Falcon m Banded Whiteface m Australian Hobby 0 Western Flyeater a Malleefowl m Redthroat a Little Button-Quail m Splendid Fairy-Wren m Spotless Crake m Rufous-crowned Emu-Wren m Black-tailed Native-hen 0 Striated Grasswren 0 Dusky Moorhen Spinifex-bird m Eurasian Coot 0 Rufous Tree-creeper p Australian Bustard 0 Red-browed Pardalote m Black-fronted Plover 0 Purple-gaped Honeyeater m Oriental Plover s Black Honeyeater 0 Red-kneed Plover s White-plumed Honeyeater 0 Red-capped Plover 0 Grey Honeyeater m Australian Dotterel m Orange Chat m Common Sandpiper 0 White-fronted Chat a Marsh Sandpiper s Magpie-lark 0 Sharp-tailed Sandpiper s White-browed Woodswallow 0 Wood Sandpiper b Little Woodswallow 0 Black-winged Stilt 0 Australian Magpie 0 Banded Stilt a Australian Crow m

82 The more mobile birds (non-passerines) were poorly represented in our samples; overall, 51 (j{ of known species were recorded, but only 24% were recorded on quadrats (Table 3). The quadrat-based strategy of survey was expected to be more effective for sampling relatively sedentary species with smaller home ranges and therefore greater densities in suitable habitat. Overall, 75170 of the passerine species previously known from the Study Area were recorded during our field work, and 61!Jo of these were recorded on quadrats (Table 3). In comparison, sampling was restricted to just 10 quadrats representing only eight of the 70 surface-types recognised from the geological maps (sce Tables I and 2). Each quadrat was sampled daily over five days during each of the three survey periods. The birds recorded on the quadrats arc listed in Table 11. Combined, the species richness of the ten quadrats did not change much from season to season; a total of 44 (33 passerine) species were recorded in summer 1979, 46 (30) in autumn 1980, and 47 (35) in winter 1981. In summer, 670 (539 passerines) individual birds were recorded on the quadrats; autumn and winter sampling yielded 896 (714) and 1061 (868) bird sightings respectively. Overhead flocks of Budgerigars, Galahs, White-fronted Honeyeaters and Masked Woodswallows accounted for 33% of these observations, and reflected the unusually high rainfall during the period of our survey. Seasonal differences could be accounted for by the number of White­ fronted Honeyeaters seen in winter (268). Quadrats were classified according to the composition of their passerine assemblages (Figure 15). The data were sufficient to show a consistent relationship between the quadrats' surface stratigraphy and their passerine composition - a separation of saline, sandy, and alluvial surfaces that is similar to the relationships noted for the other vertebrate groups. The greater mobility of the birds probably explains the different assignment of IW04, the ecotonal quadrat. There was a permanent pool of water adjacent to IW04 that attracted a variety of birds from adjacent Qpc habitats to the east and north. Despite the strong patterns in species composition, we did not find significant correlations between vertebrate richness and either floristic richness or number of vegetation strata. Such correlations were found in the Kurnalpi-Kalgoorlie Study Area (McKenzie et al. 1992), in the Nullarbor region (McKenzie and Robinson 1987) and in the Wheatbelt (Kitchener 1982, Kitchener et al. 1980). The Sandstone-Sir Samuel and Laverton-Leonora Study Areas are located in a region where ephemeral plants form a greater proportion of the plant assemblages, and where vegetations tend to be lower in stature and have fewer distinct strata, than their counterparts in the more seasonal, less arid areas to the south. Pianka (1989) has summarised some of the factors that bear on reptile richness in such regions.

REFERENCES Anon. (1993). Observations. W.A. Bird Notes 65,2-3. Austin, M.P. (1984). Problems of vegetation analysis for nature conservation. In: Survey Methods for Nature Conservation. (eds: K. Myers, CR. Margules and I. Musto) Vo!. I, pp 101-130. CSIRO Division of Water and Land Resources, Canberra. Beard, l.S. (1980). Aeolianlandforms, Western Australia. (Map at scale of 1:3,000,000 as part of Vegetation Survey of Western Australia). Vegmap Publications, Perth. Belbin, L. (1989). PATN user's guide. CSIRO Division of Wildlife and Ecology, Canberra. Belbin, L. (1991). Semi·strong Hybrid Scaling: a new ordination algorithm. 1. Veg. Sci. 2,491-496.

83 Biological Surveys Committee of Western Australia (1984). The Biological Survey of the Eastern Goldfields. Part I: Introduction and Methods. Rec. West. Aust. Mus. Supplement No. 18,1-19. Blakers, M., Davies, SJJ.F. and Reilly, P.N. (l984).The Atlas of Australian Birds. Royal Australasian Ornithologists Union. Melbourne University Press, Carlton. Buchler, E.R. (1976). A chemiluminescent tag for tracking bats and other small nocturnal animals. 1. Mammal 57, 173-176. Burbidge, A.A. and McKenzie, N.L. (1989). Patterns in the modern decline of Western Australia's vertebrate fauna: causes and conservation considerations. Biological Conservation 50, 143-198. Corben, C (1991). ANABAT ll. Bat cal/ analysis system, Version 1.1. Titley Electronics, NSW. Curry, PJ. (1979). Long-toed Stints, Sanderling and other waders at Lake Violet, Central WA in midsummer 1977-78. West. Aust. Nat. 14,109-113. Czekanowski, J. (1932). Coefficient of racial likeness, und durschschnittliche differenz. Anthrop. ANZ. 9, 227­ 249. Dell, J. and How, R.A (1988). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 5: Edjudina-Menzies Study Area. Rec. West. Aust. Mus. Supplement No. 31, 38-68. Dell, 1. and How, R.A (1992). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 6: Youanmi-Leonora Study Area. Rec. West. Aust. Mus. Supplement No. 40, 20-32. Fullard, J., Koehler, K., Surlykke, A. and McKenzie, N.L (1991). Echolocation and flight morphology of insectivorous bats in southwestern Australia. Aust. 1. Zoo!' 39,427-438. Gambold, N. and Woinarski, J.C.z. (in press). Distribution patterns of herpetofauna in monsoon rainforests of the Northern Territory, Australia. Aust. 1. Eco!. Henry-Hall, NJ. (1990). Nature conservation reserves in the Eastern Goldfields, Western Australia (southern two-thirds ofCTRC System 11). Unpublished report submitted to the EPA Red Book task force. Environmental Protection Authority: Perth, Western Australia. How, R.A., Dell, J. and Humphries, W.F. (1987). The ground vertebrate fauna of coastal areas between Busselton and Albany, Western Australia. Rec. West. Aust. Mus. 13,553-574. How, R.A. and Dell, J. (1992). Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 7: Duketon-Sir Samuel Study Area. Rec. West. Aust. Mus. Supplement No. 40, 90-103. Kitchener, DJ. (1982). Predictors of vertebrate species richness in nature reserves in the Western Australian wheatbelt. Aust. Wildl. Res. 9, 1-7. Kitchener, DJ., Chapman, A., Dell, J., Muir, B.G. and Palmer, M. (1980). Lizard assemblage and reserve size and structure in the Western Australian wheatbelt - some implications for conservation. Biological Conservation 17, 25-62. McKenzie, N.L. (1984). Biological surveys for nature conservation by the Western Australian Department of Fisheries and Wildlife: A current view. In: Survey Methods for Nature Conservation. (eds: K. Myers, CR. Margules and L Musto) VoL 2, pp 88-117. CSIRO Division of Water and Land Resources, Canberra. McKenzie, N.L and Robinson, A.C. (eds) (1987). A Biological Survey of the Nul/arbor Region, Australia. South Australian Department of Environment and Planning, Adelaide. McKenzie, N.L, Rolfe, J.K. and Carter, D. (1987). Reptiles and Amphibians. In: A Biological Survey of the Nul/arbor Region, Australia. (eds: N.L McKenzie and AC. Robinson) pp 179-210. South Australian Department of Environment and Planning, Adelaide. McKenzie, N.L and Start, AN. (1989). Structure of bat guilds in mangroves: environmental disturbance and determinism. In: Patterns in the Structure ofMammalian Communities. (eds: D.W. Morris, Z. Abramski, BJ. Fox and M.R. Willig), pp 167-178. Spec. PubL Mus. Texas Tech. Univ. No. 28. Lubbock, Texas. McKenzie, N.L, Robinson, A.C and Belbin, D.L (l99Ia). Biogeographic survey of the Nullarbor district, Australia. In: Nature Conservation: Cost Effective Biological Surveys and Data Analysis. (eds: Margules, CR. and Austin, M.P). pp 109-126. CSIRO Division of Wildlife and Ecology, Canberra.

84 McKenzie, NL, Belbln, L, Kelghery, G.J. and Kenneally, K.F (1991 b). Kimberley ramforest communities: patterns of species composition and Ilolocene biogeography. In: Kimberley Rainforests. (eds: N.L McKenzie, R.B. Johnston and PG. Kendriek) pp 423··51 Surrey Beatty and Sons, New South Wales. McKenzie, N.L, Rolfe, J,K, and Youngson, W,K. (1992), Vertebrate fauna, In: The Biological Survey of the Eastern Goldfields of Western Australia, Part 8, The Kurnalpi Kalgoorlie Study Area, Rec. West. Aust, Mu,\', Supplement No, 41,37-64, Moriarty, T.K. (1972) Birds of Wanjarri (27"35', 120°40') Emu, 72, 1-7. Pianka, E,R. (1969). Habitat specifieity, speeiation and species density in Australian desert lizards, Ecol. 50, 498-502. Pianka, E,R, (1989). Desert Lizard diversity: additional comments and some data. Am, Nat, 134, 33~··364, Pianka, H,D. and Pianka, ER, (1970) Bird censuses from desert localities in Western Australia, Emu. 70, 17·· 22. Storr, G.M. (1985). Birds of the Mid-eastern Interior of Western Australia, Rec. West. Ausl. Mus, Supplement No. 22. StOlT, G,M. (1986), Birds of the South-eastern Interior of Western Australia, Rec, West, Ausl. Mus, Supplement No. 26,

ACKNOWLEDGEMENTS We are grateful to the Curator of the Western Australian Herbarium for making available facilities generally, and to KF. Kenneally and members of the technical staff for aiding the processing of the specimens, We record our appreciation to the Director and staff of the W,A, Museum for their assistance with identification and curation of the vertebrate specimens, In particular, A, Baynes, N, Cooper, J, Henry, DJ, Kitchener, LA. Smith and G,M, Storr. Assistance with the identification of the plants was provided by TE.H. Aplin, M,I.H, Brooker, R. Carolin, RJ. Chinnock, BJ. Conn, D.A. Cooke, HJ. Eichler, J.W, Green, HJ. Hewson, S.D, Hopper, KF. Kenneally, N.S. Lander, DJ. McGillivray, TO, Mcfarlane, N,G. Marchant, B.R. Maslin, K.R. Newbey, G. Perry, B.L Rye, P,S, Short, H.R. Toelken, M, Trudgen, J.G. West, P. Weston and P.G. Wilson. R.A. How, J. Dell and KR. Newbey provided useful discussion, The Australian Bureau of Meteorology, Perth, provided the climatic data. We thank Tom Smith, Ken Cashin, Doug Watkins and Alex Harris for assistance with the prolonged and often uncomfortable field program and, in the case of Ken Cashin, for help with data compilation,

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