RECORDS OF THE WESTERN AUSTRALIAN MUSEUM 83 375–404 (2013) DOI: 10.18195/issn.0313-122x.83.2013.375-404 SUPPLEMENT

A taxonomic overview and key to the of Barrow Island, Western Australia

B. E. Heterick

Department of Environment and Agriculture, Curtin University, Bentley, Western Australia 6102, Australia. Email: [email protected]

ABSTRACT – This work characterises the (: Formicidae) fauna of Barrow Island, Western Australia, and provides a key to the workers and several unique reproductives of the 117 species recorded from the island thus far. In all, 11 of the 13 subfamilies of Western Australian ants have been recorded from Barrow Island, but Myrmeciinae and Heteroponerinae are absent. At a generic level, the fauna of the island is less rich, holding 36 of the 71 genera currently known from Western Australia. The ant fauna is characteristic of the Eremaean Botanical Province of the Pilbara, rather than that of the Carnarvon Basin from which Barrow Island is geologically derived. Ninety-three ant species (79.5% of the total on Barrow Island) are shared with the ant fauna of the Pilbara region on the adjoining mainland, but only 52 species (44.4% of the total) are shared with the ant fauna of the Carnarvon Basin. The island is very rich in unspecialised and thermophilic ant species. Five such genera, i.e., Iridomyrmex (14 spp.), Monomorium (13 spp.), Polyrhachis (12 spp.), Melophorus (10 spp.), and Camponotus (nine spp.) make up almost 50% (i.e., 49.6%) of the island’s ant fauna. Very few ants appear to be endemic to Barrow Island. The relative proportions of the two major subfamilies (Formicinae and , together comprising 61.5% of the total ant richness) are similar to the proportions found in the South-west Botanical Division for these two subfamilies (i.e., 65.9%), with Barrow Island having a slightly lower ratio of formicines to myrmicines than is found in the south-west of the state. An estimate of the total number of ant species likely to occur on Barrow Island, using the Estimate-S program (Colwell 2009), suggests that a maximum of fourteen additional species may be as yet unrecorded.

KEYWORDS: Barrow Island, ant fauna, taxonomic key, Chevron Australia

INTRODUCTION survey since 2005 (Callan et al. 2011) by Curtin University students and by employees associated Barrow Island is a 202 km2 island located some with the Gorgon natural gas project (Chevron 50 km off the north-west Australian coast, with the Australia). Among the more than 2000 terrestrial ports of Onslow and Dampier being the nearest invertebrate species collected since 2005 have been major population centres on the adjoining Australian mainland. The climate on the island is warm to hot, 117 species of ants. Accurately documenting the with the highest maxima occurring in January and of the ants collected on Barrow Island February (33.2˚C and 33.3˚C, respectively) and the has been a slow and painstaking process. This is lowest minima in July and August (17.7˚C and 17.8˚C, partially due to our still incomplete knowledge of respectively). Overall, the average annual maximum the West Australian ant fauna, and partially due is 29.1˚C and the average annual minimum is to the isolation of Perth from the main centres 22.2˚C. The mean annual rainfall is 324.6mm, most of myrmecological expertise and the major ant of it falling in the period January to May (Bureau collections on the Eastern Australian seaboard. A of Meteorology). This is the period during which short summary of the history of Western Australian cyclones or monsoonal lows affect Australia’s ant taxonomy is in order: the ants of Western northern half. In appearance the island is rather Australia have been described in a mainly piecemeal uniform with flat spinifex grasslands dominating, fashion for a century-and-a-half, with most early interspersed with termite mounds. However, more taxonomic publications resulting from collections than 227 other plants can be found on the island made by foreign researchers who came to Western (Australian Broadcasting Corporation 2011), many Australia on expeditions, e.g., W. Michaelsen and R. of them arid-adapted. Fauna includes 15 terrestrial Hartmeyer’s expedition to South-western Australia, and seven marine mammals (Bamford and Bamford E. Mjöberg’s 1910–1911 expedition to North-western 2005). Australia (the ants were described by Forel in 1907 Barrow Island has been subject to a baseline and 1915, respectively) and W. M. Wheeler, who 376 B.E. HETERICK visited Rottnest Island in 1931 (Wheeler 1934). north of the Australian mainland. Four Barrow Other significant taxonomic contributions made by Island ant species, including the two mentioned overseas researchers include those of F. Smith (1858, above, have only been collected as queens or males. 1877); C. Emery (e.g., 1895, 1898) and W. C. Crawley At the subfamily level, the fauna is taxonomically (1915, 1922). Important early Australian researchers rich, with 11 subfamilies represented out of the 22 of Western Australian Formicidae did not appear extant subfamilies currently recognized (Ward, 2007; until the 1920’s and were led by J. Clark (a number of Rabeling et al., 2008). At the generic level the fauna papers, most notably, 1924a, 1924b, 1926, 1930, 1934, is less diverse, reflecting the harsh, largely waterless 1936, 1938, 1943 and 1951) and Father J. J. McAreavey landscape. Only 36 of the 71 genera currently known (1947, 1949, 1956 and 1957). In the post WWII period, from Western Australia occur on Barrow Island. R. W. Taylor described several uncommon Western Large, important mainland groups absent from Australian ants and assisted in the revision of the Barrow Island or represented by only one species Australian bulldog ants (Taylor 1962, Taylor 1973, include Myrmecia (absent), Plagiolepis (absent), Ogata and Taylor 1991), while more mainstream taxa Prolasius (absent) Stigmacros (one species) and all of have been treated in revisionary works by Shattuck the Dacetini, apart from one Strumigenys species. and his colleagues (e.g., Shattuck 1993a; Shattuck No Myrmeciinae or Heteroponerinae are found on 1993b; Shattuck 1996; Shattuck and McMillan Barrow Island. On the other hand, genera with many 1998; Shattuck and McArthur 2002; Shattuck unspecialized or thermophilic species are well- 2007; Shattuck 2008; Shattuck 2009) and Heterick (Heterick 2001; Heterick 2003; Heterick and Shattuck represented, e.g., Iridomyrmex (14 spp.), Monomorium 2011). Heterick has also produced a handbook on (13 spp.), Polyrhachis (12 spp.), Melophorus (10 spp.), the ants of South-western Australia, with several and Camponotus (nine spp.). Together, these five nomenclatural amendments but no new species genera make up almost 50% (i.e., 49.6%) of the descriptions (Heterick 2009). Despite these efforts, Island’s ant fauna. however, the Western Australian ant fauna is still An interesting feature of the Barrow Island ants is relatively poorly known compared with that of the the relative proportions of the two major subfamilies, eastern Australian states. A name cannot confidently Formicinae and Myrmicinae. Formicinae comprise be assigned to about 38% of the morphospecies of 38.9% of the morphospecies of ants from the South- the comprehensive Western Australian holdings in west Botanical Province held in the Curtin Ant the Curtin Ant Collection, and to more than 46% Collection, and Myrmicinae comprise 27.0%. This of the taxa whose range lies outside of the South- disparity is rather less pronounced on Barrow western Botanical Division (i.e., around 133 spp.). Island, with Formicinae accounting for 31.6% of the morphospecies compared with 29.9% for CHARACTERISTICS OF THE BARROW ISLAND the Myrmicinae. On Barrow Island, a few large ANT FAUNA genera in these two groups are the most significant contributors; for the Formicinae, Polyrhachis, 76 of the 117 species of Barrow Island ants can be Camponotus and Melophorus comprise 83.8% of assigned a name, this constituting 65.0% of the ant the formicine morphospecies, and Monomorium, fauna of the island. The taxa themselves are very Meranoplus (six spp.), Pheidole (five spp.) and representative of the broader Eremaean ant fauna, Tetramorium (three spp.) comprise 77.1% of the with 93 species shared between Barrow Island and myrmicines (Table 1). the adjoining mainland. This represents 79.5% of the ants recognised from Barrow Island and 38.0% Since invertebrate sampling has taken place of the 245 Pilbara ants identified in Heterick et al. on Barrow Island annually since 2005, there are (2010). By way of contrast, only 52 ant species from sufficient data for the Island’s total ant richness to Barrow Island (i.e., 44.4% of the total recognized) are be assessed. This was done using the EstimateS included within the 243 species of ants identified by program, version 8.2 (Colwell 2009) with four Gunawardene and Majer (2004) from the southern different estimators selected. All four species Carnarvon Basin, the latter reflecting a much more accumulation curves show a strong flattening with temperate fauna. This is an interesting observation added sampling. The sampling based rarefaction in view of the fact that Barrow Island was part of curve (Mao Tau), examining a sub-sample of the Carnarvon Basin until 8000 years ago, when the pooled total species richness, produced the it was separated by rising sea levels (Eldridge et most conservative estimate of 111 species, which al. 1999). Five of the ants collected from Barrow actually slightly understates the present record Island are definitely not represented elsewhere in of 117 morphospecies. Total richness estimators the Curtin Ant Collection and, of these, four are (Chao 1 and Jack 1) predicted 131 and 127 (126.5) almost certainly unnamed (i.e., Rhytidoponera ?micans species, respectively. Bootstrapping resulted in an complex sp. JDM 1129, Discothyrea sp. JDM 1130 (a intermediate estimate of 118 species (118.3). These queen), Carebara sp. JDM 1131 and Meranoplus sp. results suggest that the ant fauna of Barrow Island, JDM 1133), while Probolomyrmex latalongus Shattuck although comprehensively sampled, may yet yield et al. (2012) (a queen) also occurs elsewhere in the up to 14 taxa that are currently unrecognised. ANTS OF BARROW ISLAND 377 Monomorium sydneyense Forel Monomorium sydneyense complex sp. JDM 101 Forel Pheidole mjobergi Pheidole sp. JDM 177 (nr variabilis Mayr) Pheidole sp. JDM 536 Pheidole sp. JDM 684 Pheidole sp. JDM 1134 Solenopsis belisarius Forel Solenopsis clarki Crawley Strumigenys sp. JDM 1230 sjostedti Forel Tetramorium spininode Bolton Tetramorium striolatum Viehmeyer Tetramorium PONERINAE Mayr Anochetus rectangularis Shattuck and Slipinska Anochetus renatae Hypoponera sp. JDM 1142 Leptogenys cf. tricosa Taylor Leptogenys sp. JDM 1128 Smith Odontomachus ruficeps F. Pachycondyla denticulata (Kirby) Pachycondyla lutea (Mayr) PROCERATIINAE ( Ƃ ) sp. JDM 1130 Discothyrea latalongus (Shattuck, Probolomyrmex and Heterick) ( Ƃ ) Gunawardene PSEUDOMYRMECINAE Smith punctulata F. Tetraponera oup sp. JDM 1175 Polyrhachis ?melanura Kohout Polyrhachis seducta Kohout Polyrhachis senilis Forel Polyrhachis sp. JDM 703 Polyrhachis sp. JDM 807 Polyrhachis sp. JDM 808 Polyrhachis sp. JDM 1009 Polyrhachis sp. JDM 1010 Polyrhachis termitoxena Wheeler Stigmacros LEPTANILLINAE Leptanilla swani Wheeler ( ƃ ) MYRMICINAE Cardiocondyla atalanta Forel Cardiocondyla nuda (Mayr) sp. JDM 1131 Carebara laeviceps chasei Forel Crematogaster sp. JDM 1132 Crematogaster Smith Meranoplus dimidiatus F. Smith Meranoplus fenestratus F. Meranoplus sp. JDM 268 Meranoplus sp. JDM 865 Meranoplus sp. JDM 889 Meranoplus sp. JDM 1133 Monomorium ‘ antipodum ’ Heterick Monomorium arenarium Monomorium disetigerum Heterick Heterick Monomorium eremophilum Monomorium euryodon Heterick Monomorium fieldi Forel Monomorium insolescens Wheeler Monomorium laeve Mayr Monomorium leae Forel Monomorium punctulatum Heterick Monomorium rubriceps gr el Shattuck and McArthur Camponotus evae zeuxis Forel Camponotus evae complex sp. JDM 1158 Camponotus fieldeae Forel Camponotus scratius Forel Camponotus simpsoni McArthur Melophorus insularis Wheeler Melophorus ludius Forel Melophorus ludius sulla For ECTATOMMINAE Rhytidoponera ? micans complex sp. JDM 1129 Rhytidoponera crassinoda (Forel) Rhytidoponera taurus (Forel) and Douglas Rhytidoponera tyloxys Brown FORMICINAE Camponotus capito Mayr Camponotus cf. evae (sp. JDM 1116) Camponotus discors Forel Camponotus donnellani Melophorus marius Forel Melophorus turneri Forel Melophorus sp. JDM 520 Melophorus sp. JDM 532 Melophorus sp. JDM 545 Melophorus sp. JDM 897 Melophorus sp. JDM 1063 Nylanderia braueri glabrior (Forel) Opisthopsis haddoni rufoniger Forel minutula (Forel) Paraparatrechina sp. JDM 916 Paraparatrechina longicornis (Latreille) Paratrechina ammonoeides Roger Polyrhachis bohemia Kohout Polyrhachis gravis Clark Polyrhachis inconspicua Emery Polyrhachis associated with Curtin University. researchers by the period 2005–2012 Island for List of ant species collected on Barrow AENICTINAE Aenictus turneri Forel AMBLYOPONINAE Amblyopone sp. indet. ( ƃ ) CERAPACHYINAE (Clark) Cerapachys brevis Cerapachys longitarsus (Mayr) Cerapachys ruficornis (Clark) Cerapachys sp. JDM 942 Cerapachys sp. JDM 1103 Cerapachys sp. JDM 1170 Sphinctomyrmex duchaussoyi (André) Arnoldius sp. JDM 433 Wheeler rottnestensis Doleromyrma Iridomyrmex agilis Forel Iridomyrmex anceps (Roger) Iridomyrmex cephaloinclinus Shattuck Iridomyrmex chasei Forel Iridomyrmex coeruleus Heterick and Shattuck Iridomyrmex discors Forel Clark Iridomyrmex dromus Iridomyrmex exsanguis Forel Iridomyrmex gibbus Heterick and Shattuck Iridomyrmex hartmeyeri Forel Iridomyrmex minor Forel Forel Iridomyrmex mjobergi Iridomyrmex sanguineus Forel Iridomyrmex tenuiceps Heterick and Shattuck Ochetellus flavipes (Kirby) Ochetellus sp. JDM 527 sp. JDM 78 sp. JDM 981 Tapinoma TABLE 1 TABLE 378 B.E. HETERICK

FIGURE 1 Presence/absence species accumulation curve illustrating estimated number of ant species on Barrow Island based on four estimators (Mao Tau ( ), Chao 1 Mean ( ) Jackknife 1 Mean ( ) and Bootstrap Mean ( )) in EstimateS version 8.2 (Colwell 2009).

KEY TO THE ANTS OF BARROW ISLAND 2. Abdominal segments III-VII with distinct constrictions between divisions so as For technical terms used in the key below, the to present an uneven outline (Figure 3) reader is referred to Heterick (2009, pp. 10–11, and (Sphinctomyrmex) ...... also pp. 198–201 (glossary))...... Sphinctomyrmex duchaussoyi André (Nb. This key recognises mainly workers, but the two queens mentioned above are also included Abdominal segments III-VII with divisions as they are the only representatives of their smoothly joined, so that the outline is even respective genera. Amblyoponinae, represented by (Figure 4) (Cerapachys) ...... 3 one unidentified male, is also included. Numbers 3. Dorsal surface of rounded onto in the case of unnamed species refer to vouchers lateral surfaces, lateral carinae absent or used in the Curtin University Ant Collection.) vestigial (Figure 5) ...... Cerapachys longitarsus (Mayr) 1. Dorsum of pygidium flattened, margins of flattened area armed laterally, posteriorly, Dorsal surface of mesosoma delimited from or both, with a series of denticles or short lateral surfaces by distinct carinae (Figure 6) spines (Figure 2); abdominal segments V – ...... 4 VII with spiracles visible in intact specimen; 4. Body concolorous red ...... 5 promesonotal suture usually completely absent (absent in all known Australian species) ...... Body black or bicoloured ...... 6 (Cerapachyinae) ...... 1 5. Ocelli present; posterior corners of head with weakly defined to strong but incomplete Dorsum of pygidium without series of dorsolateral carina curving towards eye denticles or short spines; abdominal segments (Figure 7) ...... Cerapachys sp. JDM 1103 V-VII with spiracles visible only when abdomen is distended or dissected (except for Ocelli absent; posterior corners of head Aenictinae); promesonotal suture present or without dorsolateral carina curving towards absent ...... 8 eye (Figure 8) ...... Cerapachys sp. JDM 1170 ANTS OF BARROW ISLAND 379

6. Petiole distinctly lighter coloured (yellow to 8. Waist consisting of a single distinct segment light brown) than mesosoma and post-petiole (the petiole); abdomen may be more-or-less ...... Cerapachys brevis (Clark) deeply impressed behind segment III (Figure 11) ...... 9 Petiole black, the same colour as the mesosoma and postpetiole ...... 7 Waist consisting of two distinct segments 7. Petiolar node a posteriorly rounded square in (the petiole and the postpetiole), segment III dorsal view with its posterior margin rimmed being distinctly separate from remaining by a narrow membrane; bright orange abdominal segments, which are smoothly contrasting with black postpetiole (Figure 9) rounded (Figure 12) ...... 83 ...... Cerapachys sp. JDM 942

Petiolar node rectangular, without a narrow 9. Apex of hypopygium with a circular or membrane, its posterior angles denoted by semi-circular cone (the acidopore), usually small denticles; gaster uniformly black, the projecting as a nozzle and modified to spray same as the postpetiole (Figure 10) ...... formic acid and often fringed with setae ...... Cerapachys ruficornis (Clark) (Formicinae) (Figures 13, 14) ...... 10

FIGURE 2 FIGURE 3 FIGURE 4

FIGURE 5 FIGURE 6 FIGURE 7 FIGURE 8

FIGURE 9 FIGURE 10 FIGURE 11

FIGURE 12 FIGURE 13 FIGURE 14 380 B.E. HETERICK

Apex of hypopygium lacking an acidopore, 15. Dorsum of node armed with two short but either terminating in a sting (modified distinct spines that are round in cross-section; ovipositor) or with a narrow, laterally oriented in profile, head lacking a sharp to blunt carina slit that is situated ventrally (e.g., Figure 15) between the eye and the vertex (Figure 22) ...... 46 ...... Polyrhachis sp. JDM 703

10. Antenna with 11 segments (including the Dorsum of node either unarmed or armed scape) (Stigmacros) ...... with two much abbreviated, flattened ...... Stigmacros termitoxena Wheeler denticles; head with sharp to blunt carina between the eye and the vertex (Figure 23) Antenna with 12 segments (including the ...... 16 scape) (other formicine genera) ...... 11 16. In dorsal view, propodeum strongly 11. Lower corner of mesosoma below propodeum attenuated and trapezoidal in appearance, its without an opening (to the metapleural posterior margin § 0.5 x its anterior margin gland) fringed with long setae, though a few (i.e., the metanotal groove); propodeum finely scattered setae may be present (Figure 16) ...... microreticulate-striate (Figure 24) ...... 12 ...... Polyrhachis sp. JDM 1009 Lower corner of mesosoma below propodeum In dorsal view, propodeum weakly attenuated with an opening (just above hind coxa) that is and rectangular in appearance, its posterior often fringed with long setae (Figure 17) .....32 margin ≥ 0.65 x its anterior margin; propodeum more noticeably striate (Figure 12. Upper plate of first gastral segment (first 25) ...... 17 tergite) approximately half the total length of gaster; spines or sharp angles present on 17. In rear view, propodeal dorsum separated propodeum and petiolar node in Western from propodeal declivity by a distinct carina; Australian species (Figure 18); one worker smaller species (HW ≤ 1.4 mm) (Figure 26) caste (Polyrhachis) ...... 13 ...... Polyrhachis inconspicua Emery Tergite of first gastral segment much less than half total length of gaster, spines In rear view, propodeal dorsum not separated always absent on body segments in Western from propodeal declivity by a distinct carina, Australian species; propodeal angle (if although there may be a bluntly defined angle; present) rounded (Figure 19); major and larger species (HW ≥ 1.7 mm) (Figure 27) ...... minor worker castes, at least, always present; ...... Polyrhachis sp. JDM 1010 media workers often present (Camponotus) .. 24 18. In dorsal view, pronotum as long as or longer 13. Node squamiform to semi-cuboidal and than mesonotum and propodeal dorsum directed posteriad, never with flattened combined; humeral angles dentate (Figure 28) dorsal plateau and usually armed with four (subgenus Chariomyrma) ...... 19 denticles or short spines, rarely with two short, erect parallel spines but never with two In dorsal view, pronotum much shorter elongate, recurved spines; propodeum either than mesonotum and propodeal dorsum unarmed or armed with short, upturned combined; humeral angles rounded, not flanges, denticles or short, flattened spines armed in WA species (Figure 29) (subgenus (Figure 20) (subgenus Campomyrma) ...... 14 Hagiomyrma) ...... 21

Node never squamiform, in Barrow Island 19. Lateral margins of mesonotum and species its dorsum always produced in propodeum with spinous processes (Figure the form of two elongate, recurved spines; 30) ...... Polyrhachis sp. JDM 808 propodeal angles armed with two long, acute spines (Figure 21) ...... 18 Lateral margins of mesonotum and propodeum entire (Figure 31) ...... 20 14. Mesosoma, gaster and legs with many short, bristly, erect setae ...... 20. In dorsal view, sculpture of dorsum of ...... Polyrhachis gravis Clark mesosoma easily visible and lacking obscuring pubescence ...... Mesosoma, gaster and legs glabrous ...... 15 ...... Polyrhachis senilis Forel ANTS OF BARROW ISLAND 381

FIGURE 15 FIGURE 16 FIGURE 17

FIGURE 18 FIGURE 19 FIGURE 20

FIGURE 21 FIGURE 22 FIGURE 23 FIGURE 24

FIGURE 25 FIGURE 26 FIGURE 27

FIGURE 28 FIGURE 29 FIGURE 30 FIGURE 31 382 B.E. HETERICK

In dorsal view, sculpture of dorsum of 28. In full-face view, head of major worker with mesosoma partially obscured by golden straight sides (Figure 38); first gastral plate pubescence ...... Polyrhachis sp. JDM 807 (tergite) of minor worker with one well- defined row of non-marginal erect setae ...... 21. Foreparts brick-red, gaster chocolate ...... Camponotus evae complex sp. JDM 1158 ...... Polyrhachis bohemia Kohout In full-face view, head of major worker, with Ground colour of entire body blackish ...... 22 convex sides (Figure 39); first gastral plate 22. Extremities of the petiolar spines hooked and (tergite) of minor worker with several poorly projected ventrad (Figure 32) ...... defined rows of non-marginal erect setae ...... Polyrhachis ammonoeides Roger ...... Camponotus simpsoni McArthur

Extremities of the petiolar spines recurved 29. In full-face view, anterior median portion but not hooked and projected ventrad (Figure of clypeus of both major and minor worker 33) ...... 23 produced as a squared-off projection with the corners at an angle of 90˚ to the lateral sectors 23. In dorsal view, sculpture of mesosoma very of the clypeus (Figure 40) ...... finely and evenly microreticulate; antennal ...... Camponotus fieldeae Forel scape with erect bristles ...... Polyrhachis seducta Kohout In full-face view, anterior median portion of clypeus not so produced, and usually In dorsal view, sculpture of mesosoma finely projecting as a weak convexity, the anterior longitudinally striate, the striae joined by median angles (if present) oblique (Figure 41) irregular cross ribs; antennal scape lacking ...... 30 erect bristles ...... Polyrhachis ?melanura Kohout 30. In profile, mesonotum and propodeum 24. Mentum of major and minor worker with of major and minor workers elongate and elongate, J-shaped setae near its posterior undulant, saddle-shaped in the minor worker margin (Figure 34) ...... (Figure 42) ...... Camponotus capito Mayr Camponotus donnellani Shattuck and McArthur In profile, mesonotum and propodeum Without elongate, J-shaped setae on posterior high in major and minor worker, gradually margin of mentum (Figure 35) ...... 25 curving into a steep propodeal declivity (Figure 43) ...... 31 25. Major and minor workers with five mandibular teeth; black or very dark brown 31. Species larger (HW major worker ≥ 2.5mm, ants with a high, truncate mesosoma and a HW minor worker ≥ 0.9mm); paired, erect steeply declivitous propodeal declivity...... 26 pronotal setae in minor worker placed in centre of pronotum (Figure 44) ...... Major and minor workers with maximum of ...... Camponotus discors Forel six or more teeth; if dentition ambiguous due to abrasion of small denticles, and with high Species smaller (HW major worker ≤ 1.4mm, mesosoma (a few C. scratius minor workers), HW minor worker ≤ 0.6mm); paired, erect then paler in colour ...... 29 pronotal setae in minor worker placed near to promesonotal suture (Figure 45) ...... 26. Body and legs notably bristly, the erect setae ...... Camponotus scratius Forel on the legs raised at an angle of ≈ 45˚ in both major and minor workers (Figure 36) ...... 32. Eyes very large and placed on posterior ...... Camponotus cf. evae (sp. JDM 1116) angles of head capsule (Figure 46) ...... Body and legs less hairy, the erect setae on the ...... Opisthopsis haddoni rufoniger Forel legs appressed or raised at an angle of ≈ 15˚ or Eyes of moderate size and placed at sides or less (Figure 37) ...... 27 front of head capsule, but not on posterior 27. A few erect setae visible on the venter of the angles (Figure 47) ...... 33 head capsule in major and minor workers ...... Camponotus evae zeuxis Forel 33. Propodeal spiracle slit or comma-like (Figure 48); clypeus and underside of head usually Erect setae completely lacking on the venter of with profuse, long, curved setae (Figure 49); the head capsule in major and minor workers polymorphic with major, media and minor ...... 28 workers present (Melophorus) ...... 34 ANTS OF BARROW ISLAND 383

FIGURE 32 FIGURE 33 FIGURE 34 FIGURE 35

FIGURE 36 FIGURE 37 FIGURE 38 FIGURE 39

FIGURE 40 FIGURE 41 FIGURE 42

FIGURE 43 FIGURE 44 FIGURE 45

FIGURE 46 FIGURE 47 FIGURE 48 FIGURE 49 384 B.E. HETERICK

Propodeal spiracle oval or round (Figures 50); 37. Mesosoma with metanotum apparently clypeus and underside of head and mandibles developed and confluent with mesonotum, with few or no long, curved setae (Figure 51); often extending over the propodeum; monomorphic species ...... 43 metanotal groove obsolete, its position indicated only by a superficial, transverse 34. In full-face view, anterior clypeal margin in all worker castes convex, apron-like and furrow; propodeum reduced in size and covering whole or part of the retracted wedge-shaped, with narrow end of wedge mandible, the medial clypeal sector often often under fold of metanotum; metathoracic produced so that it is protrusive when seen spiracle lateral and situated within metanotal in profile; clypeal psammophore frequently sector (Figure 62) ...... with coarse and well-separated ammochaetae, ...... Melophorus sp. JDM 1063 these always placed on or just above anterior margin (Figure 52); in profile, propodeum Worker mesosoma of normal appearance, elongate and oblique or broadly rounded with metathoracic spiracle situated on or near (Figure 53) ...... 35 dorsum of mesosoma (Figure 63) ...... 38 In full-face view, anterior clypeal margin in all worker castes variable, but not covering 38. Gaster with curved erect, semi-erect setae whole or part of the retracted mandible, the and a few decumbent setae only, genuine medial clypeal sector not narrowly protrusive, appressed setae lacking; body strongly although it may be broadly protuberant; sculptured and hirsute, antennal scapes and clypeal psammophore often placed along the midpoint of the clypeus or even above it legs with whorls of many fine, straight setae (Figure 54); in profile, propodeum typically ...... Melophorus sp. JDM 532 truncate or narrowly rounded (Figure 55) ...... 37 If legs and antennal scapes with whorls of setae, then gaster with at least well-spaced 35. Mesonotum and propodeum of minor worker appressed setae between the longer, erect or confluent, metanotal groove completely semi-erect pilosity ...... 39 lacking (major worker unknown) (Figure 56) ...... Melophorus sp. JDM 897 39. Tibiae and antennal scape matt, strongly Mesonotum and propodeum of minor worker microreticulate; short, erect setae present on not confluent; metanotal groove usually antennal scape and metatibia and usually well-developed, but if not then a furrow that in whorls, length of longest setae < greatest represents the metanotal groove is present width of tibia; gaster of minor worker (Figure 57) ...... 36 strongly pubescent ...... 36. In profile, mesosoma of minor worker ...... Melophorus sp. JDM 520 with a compact appearance, its dorsal outline describing a pronounced arc due to Tibiae and antennal scape lacking erect shape of the mesonotum and mesopleuron pilosity, moderately to strongly shining and (mesosternal outline and dorsum of smooth or with superficial microreticulation; mesonotum strongly convergent anteriorly) gaster of minor worker without pubescence, (Figure 58); in profile, clypeus of all workers often glabrous or nearly so ...... 40 gently recurved and produced as a small ledge over basal sector of mandibles (Figure 59) ...... Melophorus insularis Wheeler 40. In profile, head of minor worker not dorsoventrally compressed, its side 0.7 x as In profile, mesosoma of minor worker with wide as long ≥ (Figure 64); in full-face view, elongate gracile appearance, its dorsal outline eye set slightly above midpoint of gena; straight or describing a weak arc (mesosternal clypeal psammophore at about midpoint of outline and dorsum of mesonotum weakly convergent to subparallel anteriorly (Figure clypeus; major worker with multiple preapical 60)); in profile, clypeus of all workers metatibial spurs; larger species (HW of major produced as a flange that projects anteriad worker 1.30 mm >); erect marginal setae (and well past basal sector of mandibles (Figure 61) often non-marginal setae) present on gaster in ...... Melophorus sp. JDM 545 minor worker ...... 41 ANTS OF BARROW ISLAND 385

FIGURE 50 FIGURE 51 FIGURE 52 FIGURE 53

FIGURE 54 FIGURE 55 FIGURE 56 FIGURE 57

FIGURE 58 FIGURE 59 FIGURE 60

FIGURE 61 FIGURE 62 FIGURE 63 FIGURE 64 386 B.E. HETERICK

In profile, head of minor worker Mandible armed with five teeth; antennal dorsoventrally compressed, its side as little scape lacking erect setae ...... 44 as 0.5 x as wide as long (Figure 65); in full- 44. Propodeum lacking erect setae (Figure 68); face view, eyes set well above midpoint of erect setae scattered over head capsule; gena; clypeal psammophore on or just above femora and tibia with a few large, erect setae anterior clypeal margin in minor workers, (Paratrechina) ...... placed just above anterior clypeal margin to ...... Paratrechina longicornis (Latreille) midway between anterior clypeal margin and midpoint of clypeus in major workers; major One pair of erect setae present on propodeum worker with just one preapical metatibial (Figure 69): four erect setae on posterior spur; smaller species (HW of major worker ≤ margin of vertex, with two rows of paired erect setae extending from posterior margin 1.20 mm); erect marginal and non-marginal of vertex to posterior clypeal margin; setae lacking on gaster in minor worker femora and tibiae lacking large erect setae ...... 42 (Paraparatrechina) ...... 45

41. Colour of ant pale, depigmented yellow 45. Eye elliptical, moderate in size (eye length to bright orange-yellow in minor worker; < 0.3 x head length (Figure 70); brownish- concolorous orange-yellow or with darker yellow ...... Paraparatrechina minutula (Forel) gaster in major worker; in full-face view, colour of head capsule often two-toned, with Eye slightly asymmetrical, larger (eye length orange-yellow above the anterior margin of ≈ 0.3 x head length) (Figure 71); depigmented the eye, and beige below the eye ...... yellow ...... Melophorus ludius sulla Forel Paraparatrechina minutula group sp. JDM 916

Colour various shades of brown, reddish 46. Apex of hypopygium with a narrow, laterally brown or deep orange, concolorous or oriented slit that is situated ventrally (Figure bicoloured with gaster always darker (often 72); abdomen without an impression between dark brown or black), but never as above ...... the third and fourth abdominal segments; ...... Melophorus turneri Forel gastral cuticle often soft, flexible and easily collapsed (Dolichoderinae) ...... 47 42. In full-face view, major worker with posterior clypeal margin not arched or falling away Apex of hypopygium with a sting (Figure 73); between antennal insertion and tentorial abdomen may be deeply impressed between pit (Figure 66); anterior clypeal margin of third and fourth abdominal segments; cuticle major worker straight and never protuberant; stronger, less flexible and not normally major worker mesosoma with up to a dozen collapsible ...... 69 fine, flexuous erect setae; minor worker 47. Petiole without a distinct node (Figure 74); mesosoma glabrous, glossy, with superficial posterior margin of clypeus a broad, even arc sculpture only; colour pale yellowish to (Tapinoma) ...... 48 brown, concolorous or with darker gaster ...... Melophorus ludius Forel Petiole usually with a distinct node (Figure 75); if node weak, then posterior margin In full-face view, major worker with posterior of clypeus elliptical or forming a shallow clypeal margin arched and falling away rectangle with median sector often more-or- between antennal insertion and tentorial pit less straight ...... 49 (Figure 67); anterior clypeal margin of major worker weakly convex or with anteromedial 48. Eye large, eye length ≈ 1/3 length of head dimple indicated; major worker mesosoma capsule (Figure 76) ...... with a few short, bristly or slightly clavate ...... Tapinoma sp. JDM 981 (but not flexuous) erect setae; minor worker mesosoma glabrous with some shagreenate Eye smaller, eye length ≤ ¼ length of head sculpture and matt to moderately shining; capsule (Figure 77) ...... Tapinoma sp. JDM 78 bicoloured orange-and-brown to brown with darker gaster ...... Melophorus marius Forel 49. Palps very short (PF 2,2) (Figure 78); eyes small (about 50 facets); clypeus with several 43. Mandible armed with six or seven teeth; to many downwardly curved setae, which are antennal scape with erect setae (Nylanderia) about the same length as the closed mandible ...... Nylanderia braueri glabrior (Forel) (Arnoldius) ...... Arnoldius sp. JDM 433 ANTS OF BARROW ISLAND 387

FIGURE 65 FIGURE 66 FIGURE 67 FIGURE 68

FIGURE 69 FIGURE 70 FIGURE 71

FIGURE 72 FIGURE 73 FIGURE 74

FIGURE 75 FIGURE 76 FIGURE 77 FIGURE 78 388 B.E. HETERICK

Palps longer (PF 6,4) (Figure 79); eyes 54. Acute anteromedial clypeal prominence generally larger; clypeus with several to present as a conspicuous triangle (Figure 90); many short, occasionally curved setae that are erect setae present on antennal scapes and shorter than the closed mandible ...... 50 hind femora, and often a few on sides of head; frontal carinae distinctly sinuate with small flanges on carinae often tending to angulate 50. Declivitous face of propodeum concave; medially; metathoracic spiracles prominent gaster often shining with purple or blue (medium-large to large ants, often with broad, iridescence (Figure 80); nearly always triangular heads and with weak pink or associated with wood or manmade structures purplish iridescence on body and legs) ...... (Ochetellus) ...... 51 ...... 55

Declivitous face of propodeum never concave, Anteromedial clypeal prominence not a usually rounded (e.g., Figure 81); if iridescent, conspicuous triangle (either a small blunt then iridescence extended to mesosoma and projection, an indistinct undulation of the head; northern Western Australian species cuticle or completely absent) (Figure 91) nest terrestrially ...... 52 or erect setae absent from one or more of antennal scapes, sides of head and hind femora; frontal carinae rarely distinctly 51. In profile, propodeum vertically rectangular sinuate ...... 56 (propodeal length ≤ 0.40 x propodeal width); in dorsal view, propodeum terminating in a 55. Antennal scapes short (SL < 1.30mm); in full- tapered flange (Figures 82a, b) ...... face view, anteromedial clypeal prominence ...... Ochetellus flavipes (Kirby) shorter, not extending beyond lat eral lobes of anterior clypeal margin; erect hairs on head and body fine and pale in colour……..... In profile, propodeum square, about as Iridomyrmex discors Forel long as high; in dorsal view, propodeum terminating in a squared-off flange (Figures Antennal scapes long (SL > 1.35mm); in full- 83a, b) ...... Ochetellus sp. JDM 527 face view, anteromedial clypeal prominence longer, extending beyond lateral lobes of 52. Mesosoma consisting of three compact anterior clypeal margin; erect hairs on head segments, the two thoracic segments higher and body typically bristly and dark in colour . than long, the propodeum obliquely flattened ...... Iridomyrmex sanguineus Forel (Figure 84); node indistinct, barely rising above the articulation of the peduncle with 56. Semi-erect to erect setae present on antennal propodeum; small to minute ants (1–1.5 mm) scapes and hind tibiae and, often, sides of (Doleromyrma) ...... head (a few hirsute populations of I. chasei that may occur on Barrow Island) ...... Doleromyrma rottnestensis Wheeler ...... Iridomyrmex chasei Forel (pt.)

Mesosoma less compact, the propodeum Semi-erect to erect setae absent from antennal generally rounded or roundly cuboidal (e.g., scapes, or hind tibiae, or both parts ...... 57 Figure 85); petiolar node distinct; generally larger species (2–5 mm) (Iridomyrmex) ...... 53 57. Hind tibiae with distinct erect and/or semi-erect setae in addition to appressed or 53. In profile, petiolar node thick, very elongate decumbent setae (may be sparse) ...... 58 and strongly inclined anteriad, the anterior face very short or even virtually absent Hind tibiae without erect/sub-erect setae (Figure 86); in full-face view, frontal carinae ...... 59 strongly concave (Figure 87) ...... Iridomyrmex cephaloinclinus Shattuck 58. In full-face view, erect setae confined to posterior margin of head (Figure 92); erect setae on hind tibiae sparse, often confined In profile, petiolar node not as above (Figure to one or two (a few hirsute populations of I. 88); frontal carinae weakly concave, straight anceps that may occur on Barrow Island) ...... or weakly convex (Figure 89) ...... 54 ...... Iridomyrmex anceps (Roger) (pt.) ANTS OF BARROW ISLAND 389

FIGURE 79 FIGURE 80 FIGURE 81

AB AB

FIGURE 82 FIGURE 83

FIGURE 84 FIGURE 85

FIGURE 86 FIGURE 87 FIGURE 88

FIGURE 89 FIGURE 90 FIGURE 91 FIGURE 92 390 B.E. HETERICK

In full-face view, erect setae extending along Mesosoma with six ≥ erect setae, seta often lateral margins of head to at least the eyes numerous; in profile, propodeum may be (Figure 93); typically, erect setae on hind smoothly rounded rather than protuberant; tibia moderately plentiful to abundant and posterior margin of head generally slightly extending along length of tibia (a few hirsute convex or planar, but may be very weakly populations of I. minor that may occur on concave ...... 63 Barrow Island) ...... Iridomyrmex minor Forel (pt.) 63. Propodeal dorsum rounding on to declivitous face through a protuberance (Figure 98) ...... 59. Head in full-face view extremely narrow (CI ...... Iridomyrmex exsanguis Forel 64-70); in dorsal view, pronotum very weakly tapering towards its anterior margin; eye Propodeal dorsum evenly rounded on to large (EL > 0.24mm, EW > 0.19 mm, EI > 38, declivitous face without a protuberance eye width greater than or equal to 3 x greatest (Figure 99) ...... Iridomyrmex dromus Clark diameter of antennal scape) ...... Iridomyrmex tenuiceps Heterick and Shattuck 64. Antennal scape surpassing posterior margin of head capsule by at least 3 x its width and Without above combination of three often 6 x its width; vertex of head capsule characters (in I. agilis, which is similar, the eye slightly convex to very weakly concave; is smaller) ...... 60 mesosoma gracile with propodeum that is an elongate curve in profile ...... 65 60. Head in full-face view elongate (CI < Antennal scape surpassing posterior margin 83), upper vertex rather broad above eyes of head capsule by 2 x its width ≤ ; vertex of (Figure 94); in the same view, eyes placed head may be deeply concave; mesosoma more at about midpoint of head; margin of compact, in profile often with truncate and clypeus produced as a sharp, narrowly triangular tooth; hind femur very long, raised propodeum ...... 66 length greater than or equal to mesosoma 65. Concolorous brown to dark brown, often ...... Iridomyrmex agilis Forel with coppery reflections; hairs on pronotum and mesonotum short and bristly, their If head elongate, length of hind femur less length not exceeding greatest diameter of than length of mesosoma or, in full-face view, the eye; length of hind femur variable, but eyes placed above midpoint of head and usually less than 0.90 × length of mesosoma upper vertex narrow compared with region ...... Iridomyrmex anceps (Roger) (pt.) below eyes (e.g., Figure 95), and anteromedial clypeal margin not produced as a sharp, Not concolorous brown (generally, gaster narrowly triangular tooth ...... 61 darker than body, and head and foreparts often with varying degrees of reddish 61. Eye with distinctly protuberant anteromedial or orange coloration); length of hairs on margin, and a posterior lobe that is narrower pronotum and mesonotum often exceeding than its anterior lobe (Figure 96); colour of greatest diameter of the eye; length of hind ant most commonly depigmented or tawny femur greater than or equal to 0.90 × length of yellow, but never iridescent ...... 62 mesosoma ...... Iridomyrmex minor Forel (pt.)

Eye with, at most, a slightly protuberant 66. In full-face view, posterior margin of head anteromedial margin, and a posterior lobe broadly concave, posterolateral corners that is not narrower than its anterior lobe broadly angulate (Figure 100); in profile, (Figure 97); ant shades of reddish-brown, anterior pronotum humped, arising steeply brown or black (bicoloured or concolorous), at angle of ≈ 60˚; propodeum narrowly pro- and may have iridescence ...... 64 tuberant (Figure 101); head and mesosoma without iridescence; if gaster with blue-green 62. Erect hairs short, pronotal and mesonotal reflections then ground colour of gaster black hairs six <, often mesosoma glabrous; in and foreparts bright orange with or without profile, propodeum strongly protuberant; some brown infuscation, gaster usually in full-face view, posterior margin of head with coppery reflections only; pronotum weakly to strongly concave ...... usually with 10 > erect setae, never glabrous ...... Iridomyrmex hartmeyeri Forel ...... Iridomyrmex chasei Forel (pt.) ANTS OF BARROW ISLAND 391

In full-face view, posterior margin of head 67. In profile, ant very compact; anterior usually planar (Figure 102), but if broadly pronotum humped, arising at angle of concave and posterolateral corners broadly ≈ 45º and descending as a steep and angulate, then ant blackish or dark brown symmetrical curve (Figure 104); propodeum with faint to strong blue-green, yellow-green narrowly protuberant and truncate with a or pinkish iridescence; anterior pronotum not planar dorsum; head and mesosoma never so steeply humped, arising at 45˚ ≤ , and its iridescent; concolorous plain brown species dorsum may be glabrous (Figure 103) ...... 67 ...... Iridomyrmex gibbus Heterick and Shattuck

FIGURE 93 FIGURE 94 FIGURE 95

FIGURE 96 FIGURE 97 FIGURE 98

FIGURE 99 FIGURE 100 FIGURE 101

FIGURE 102 FIGURE 103 FIGURE 104 392 B.E. HETERICK

In profile, ant less compact; anterior pronotum Hind tibial spur present and distinct; apex of usually arising as a smooth curve at angle petiolar node planar or tapered and lacking ≤ 30˚ (Figure 105); in profile, propodeum a process or point directed posteriad (e.g., often broadly rounded, but if rectangular Figure 113) ...... 73 then its dorsum rounded in Barrow Island populations; head and mesosoma usually 73. In profile, petiolar node thick and cuboidal or with some iridescence, either bluish, yellow- sub-cuboidal (Figure 114) ...... green or pinkish (I. coeruleus) or coppery (I...... Rhytidoponera crassinoda (Forel) mjobergi) ...... 68 In profile, petiolar node thin and tapered 68. Dorsum of mesosoma with short, bristly towards its apex (Figure 115) ...... whitish setae; iridescence distinct and bluish Rhytidoponera ?micans complex sp. JDM 1129 in Barrow Island populations ...... Iridomyrmex coeruleus Heterick and Shattuck 74. Promesonotal suture either completely absent or present and reduced and fully fused, so Dorsum of mesosoma either glabrous or pronotum and mesonotum are incapable of with a few short, dark, bristly setae; weak, independent movement (Figure 116); antennal coppery iridescence only in Barrow Island sockets mostly to completely exposed (Figure populations…..Iridomyrmex mjobergi Forel 117) (Proceratiinae) ...... 75

69. Petiole broadly articulated to abdominal Promesonotal suture fully developed, so segment III; dentiform clypeal setae present pronotum and mesonotum capable of (Figure 106) (Amblyoponinae) ...... independent movement (Figure 118); antennal ...... Amblyopone sp. indet. (a single male ant) sockets covered by developed frontal lobes (Figure 119) (Ponerinae) ...... 76 Petiole with distinctly descending posterior face; dentiform clypeal setae absent (Figure 75. Second gastral tergite strongly arched so that 107) ...... 70 succeeding segments are ventral and oriented anteriad towards the head end of the ant 70. In profile, metapleural gland orifice a (Figure 120) (Discothyrea) ...... longitudinal to oblique curved slit or crescent, ...... Discothyrea sp. JDM 1130 (a single queen) directed upward by a strip of cuticle (Figure 108) (Ectatomminae) ...... 71 Second gastral tergite only very weakly arched, successive segments oriented In profile, metapleural gland orifice posteriad (Figure 121) (Probolomyrmex) ...... elliptical to circular and opening laterally or Probolomyrmex latalongus (Shattuck, posteriorly, not bounded by strip of cuticle Gunawardene and Heterick) (a single queen) that directs orifice upward (Figure 109) ...... 74 76. Mandibles long and linear, inserted in central 71. In full-face view, angles of vertex of head anterior margin of head (Figure 122) ...... 77 capsule produced as distinct, raised denticles (Figure 110) ...... Rhytidoponera taurus (Forel) Mandibles triangular or elongate, curved, inserted at sides of head (Figure 123) ...... 79 In full-face view, angles of vertex of head capsule more-or-less rounded angles (Figure 77. Top of head with V-shaped lines converging 111) ...... 72 to form a groove on upper front of head (Figure 124) (Odontomachus) ...... 72. Hind tibial spur highly reduced and difficult ...... Odontomachus ruficeps F. Smith to distinguish from surrounding spines; apex of petiolar node terminating in a sharp spur Top of head without V-shaped lines and (usually) or a dull point (rarely) directed with broad, uninterrupted curved ridge; posteriad (Figure 112) ...... weak groove present or absent (Figure 125) ...... Rhytidoponera tyloxys Brown and Douglas (Anochetus) ...... 78 ANTS OF BARROW ISLAND 393

FIGURE 105 FIGURE 106 FIGURE 107

FIGURE 108 FIGURE 109 FIGURE 110

FIGURE 111 FIGURE 112 FIGURE 113 FIGURE 114

FIGURE 115 FIGURE 116 FIGURE 117 FIGURE 118

FIGURE 119 FIGURE 120 FIGURE 121

FIGURE 122 FIGURE 123 FIGURE 124 FIGURE 125 394 B.E. HETERICK

78. Entire pronotum and sides of propodeum 84. Pronotum and mesonotum fused to form one smooth and shining; dorsal surface of segment; antennae 10-segmented (Figure 132); propodeum with weak transverse striations length 3mm > (Aenictinae: Aenictus) ...... (nearly absent in some specimens); setae on ...... Aenictus turneri Forel dorsal surface of propodeum very short, scattered and appressed ...... Joint between pronotum and mesonotum ...... Anochetus renatae Shattuck flexible; antennae 12-segmented (Figure 133); length 2.5 mm < (Leptanillinae: Leptanilla) ..... Pronotum partially to completely sculptured; Leptanilla swani Wheeler (males only collected sides of propodeum with coarse striations, on Barrow Island) dorsal surface coarsely sculptured with combination of irregular rugosity 85. Joint between pronotum and mesonotum and striations; setae on dorsal surface of flexible (Figure 134): hind tibiae with propodeum longer, erect or semi-erect ...... pectinate spurs; tarsal claws toothed (Figure ...... Anochetus rectangularis Mayr 135) (Pseudomyrmecinae: Tetraponera) ...... Tetraponera punctulata F. Smith 79. Pretarsal claws of hind leg equipped with one or more teeth on the inner curvature, and Pronotum and mesonotum fused to form usually pectinate: clypeus produced anteriad one segment (the promesonotum) (Figure as an acute, V-shaped projection (Figure 126) 136); hind tibiae with at most a simple spur, (Leptogenys) ...... 80 but this may be lacking; tarsal claws simple (Figure 137) (Myrmicinae) ...... 86 Pretarsal claws of hind leg simple; clypeus straight or broadly convex; not produced 86. Distinctive ant with triangular, deeply emarginate head; antennal segments ≤ six; anteriad as an acute, V-shaped projection mandible curved, elongate and armed at (Figure 127) ...... 81 the tip with intersecting spikes (Figure 138) 80. In full-face view, mandibles curved, elongate; (Strumigenys) ...... Strumigenys sp. JDM 1230 eyes of normal appearance; cuticle of head, Ant not as above; head more rounded (Figure mesosoma and petiolar node with many 139); antennal segments ≥ nine; mandible shallow foveae; dark brown to blackish ants triangular ...... 87 ...... Leptogenys sp. JDM 1128 87. Antenna with nine segments; dorsum of In full-face view, mandibles narrowly anterior mesosoma flattened and projecting to triangular, their blades parallel; eyes vestigial; form a shield, often with regular protruding cuticle of head, mesosoma and petiolar node edges and translucent ‘windows’ between smooth and glossy; orange ants ...... these edges (Figure 140) (Meranoplus) ...... 88 ...... Leptogenys cf. tricosa Taylor Antenna with 10 or more segments; dorsum 81. Tibia of hind leg with a single large pectinate of mesosoma never forming a shield as above spur (Figure 128) (Hypoponera) ...... 93 ...... Hypoponera sp. JDM 1142 88. In full-face view, clypeus strongly incurved, Tibia of hind leg with both a single large weakly tapered anteriad, emarginate in pectinate spur and a smaller, simple spur appearance and extended only slightly (Figure 129) (Pachycondyla) ...... 82 beyond the apices of the antennal lobes; 82. Larger species (HW ≥ 2 mm); heavily antennal lobes broad, often hiding most of the sculptured ...... Pachycondyla denticulata (Kirby) eye (Figure 141) ...... 89

Smaller species (HW ≈ 1 mm); at most, weakly In full-face view, clypeus weakly incurved, sculptured ...... Pachycondyla lutea (Mayr) moderately to strongly tapered anteriad with a straight anteromedial margin and extended 83. Eyes absent or represented by a single facet; well beyond the apices of the antennal lobes; frontal lobes absent, so that the antennal antennal lobes narrower, so eye can often be insertions are completely exposed (Figure clearly seen (Figure 142)...... 90 130) ...... 84 89. In full-face view, sculpture of head capsule Eyes normally present, but if absent, frontal with vestigial, minute, dense longitudinal lobes expanded so that the latter cover all or striae that are almost invisible; in profile, part of the antennal insertions (Figure 131) petiolar node subcuboidal, its dorsum planar ...... 85 (Figure 143)...... Meranoplus fenestratus F. Smith ANTS OF BARROW ISLAND 395

FIGURE 126 FIGURE 127 FIGURE 128 FIGURE 129

FIGURE 130 FIGURE 131 FIGURE 132

FIGURE 133 FIGURE 134 FIGURE 135

FIGURE 136 FIGURE 137 FIGURE 138 FIGURE 139

FIGURE 140 FIGURE 141 FIGURE 142 FIGURE 143 396 B.E. HETERICK

In full-face view, sculpture of head capsule 95. First and second antennal segments much consisting of distinct, evenly spaced longer than remaining segments and forming longitudinal striae; in profile petiolar node a distinct two-segmented club (Figure 155) narrow and tapering, its dorsum rounded ...... 96 (Figure 144)...... Meranoplus sp. JDM 268 Antennae either without a distinct club or 90. In dorsal view, promesonotal shield without with a three-segmented club (Figure 156) spines or a flange on its posterior margin ...... 98 (Figure 145) ...... Meranoplus dimidiatus F. Smith 96. Rear face of propodeum with flanges (Figure In dorsal view, promesonotal shield with 157); clypeus with a pair of setae that straddle spines on its posterior margin joined by at the midpoint of the anterior clypeal margin least a small flange (Figure 146) ...... 91 (Figure 158); strongly dimorphic, major workers with a pair of short horns on the 91. In rear view, posterior face of petiolar node vertex of the head capsule in some Eastern with a series of v-shaped striae contained states species (Carebara) (a single minor within each in a nested pattern; postpetiole worker) ...... Carebara sp. JDM 1131 also striate (Figure 147) ...... Rear face of propodeum rounded, never with ...... Meranoplus sp. JDM 865 teeth, spines or flanges (Figure 159); midpoint In rear view, at least dorsum of posterior of anterior clypeal margin with a single seta face of petiolar node foveate; postpetiole also (Figure 160); WA species weakly polymorphic foveate (Figure 148) ...... 92 (Solenopsis) ...... 97 97. Eye absent or represented by a minute, 92. In dorsal view, posterolateral spines of pigmented speck (Figure 161) ...... promesonotal shield the most prominent, the ...... Solenopsis belisarius Forel posterior angles represented by narrower and usually shorter spines (Figure 149) ...... Eye small but distinct (Figure 162) ...... Meranoplus sp. JDM 889 ...... Solenopsis clarki Crawley

In dorsal view, spines at the posterior angles 98. Viewed from front, area of clypeus below of promesonotal shield the most prominent, antennal sockets raised into a sharp ridge the posterolateral spines broader but much (Figure 163); tip of sting with a triangular shorter than the former (Figure 150) ...... or club-like appendage projecting upwards ...... Meranoplus sp. JDM 1133 from the shaft (Figure 164); propodeal angle usually a pair of stout spines, sometimes 93. Postpetiole attached to upper surface of flanges (Tetramorium) ...... 99 gaster, which is heart-shaped when seen from above; petiole flattened; viewed from above, Viewed from front, area of clypeus below postpetiole often distinctively bilobed (Figure antennal sockets smooth or a dull ridge 151) (Crematogaster) ...... 94 (Figure 165); tip of sting thin and pointed, occasionally slightly flattened, but without Postpetiole attached to the front of the gaster, appendage; propodeal angle often absent or which is not distinctively heart-shaped; with pair of protuberances only (Figure 166) petiole usually with a node, not flattened; ...... 101 postpetiole not bilobed as above (e.g., Figure 99. Viewed in profile, petiolar node produced as 152) ...... 95 a spur directed posteriad (Figure 167); lateral 94. Propodeum flattened and all sectors on margins of first gastral tergite flattened to the same plane (except for a narrow strip form flanges ...... Tetramorium spininode Bolton behind the metanotal groove); anterior lateral Viewed in profile, petiolar node cuboidal and propodeal carinae present, extending from not produced as a spur directed posteriad metanotal groove to propodeal angles (Figure (Figure 168); lateral margins of first gastral 153) (subgenus Orthocrema) ...... tergite rounded towards their junction with ...... Crematogaster sp. JDM 1132 first ventral plate of gaster (sternite) ...... 100 Anterior sector of propodeum not flattened, 100. Viewed in profile, gross sculpture of often convex, not on same plane as posterior mesosoma consisting almost exclusively sector; anterior lateral propodeal carinae of parallel striae, the cuticle between them always absent (Figure 154) ...... deeply incised (Figure 169); larger species ...... Crematogaster laeviceps chasei Forel (HW 1.20 mm >) ...... Tetramorium sjostedti Forel ANTS OF BARROW ISLAND 397

FIGURE 144 FIGURE 145 FIGURE 146 FIGURE 147

FIGURE 148 FIGURE 149 FIGURE 150 FIGURE 151

FIGURE 152 FIGURE 153 FIGURE 154 FIGURE 155

FIGURE 156 FIGURE 157 FIGURE 158 FIGURE 159 FIGURE 160

FIGURE 161 FIGURE 162 FIGURE 163 FIGURE 164 FIGURE 165

FIGURE 166 FIGURE 167 FIGURE 168 FIGURE 169 398 B.E. HETERICK

Viewed in profile, gross sculpture 106. Propodeum more-or-less rounded, with of mesosoma consisting of a mixture of small, inconspicuous metapleural lobes parallel striae, costulate sculpture and (Figure 177); propodeal and mesopleural microreticulation, the cuticle between the sculpture never shagreenate-punctate, usually gross sculpture not markedly incised (Figure absent, if present, confined to a few striae, 170); smaller species (HW 1.10 mm <) ...... particularly around the katepisternum ...... 107 ...... Tetramorium striolatum Viehmeyer Propodeum distinctly cuboidal, or with 101. Central anterior margin of clypeus with a propodeal lobes lamellate and extending to single seta, which is often surrounded by near propodeum; propodeal and mesopleural paired setae (Figure 171); monomorphic sculpture may be shagreenate-punctate or weakly polymorphic (i.e., exhibiting (Figure 178)...... 108 monophasic allometry) except for Monomorium euryodon ...... 102 107. Yellow ants; propodeum relatively elongate ...... Monomorium laeve Mayr Central anterior margin of clypeus with paired setae or undifferentiated setae (Figure Brown ants; propodeum compact and 172); strongly dimorphic (Pheidole) ...... 116 rounded ...... Monomorium fieldi Forel

102. PF 5,3; clypeus plate-like, projecting and 108. Propodeum smooth and shining with recessed posteriorly towards tentorial pits, only vestigial striae; mandible with three (indented points of attachment of internal distinct teeth; eye rather small (approximately muscles), clypeus not bicarinate (Figure 173) width of antennal scape); propodeum with (Cardiocondyla) ...... 103 declivitous face long and oblique, carinate at sides and sometimes with small lamellae PF 2,3, 2,2 or 1,2; clypeus not plate-like at propodeal angle; anterior clypeal margin or recessed posteriorly towards tentorial rounded; long, erect and suberect setae absent pits, clypeus often bicarinate (Figure 174) from trunk; colour tawny ...... (Monomorium) ...... 104 ...... Monomorium arenarium Heterick

103. Head and mesosoma of worker matt or Propodeum shagreenate or otherwise weakly shining, uniformly microreticulate; sculptured or colour bright yellow; mandible in rear view, rows of appressed gastral setae with four teeth and denticles; clypeus usually longer, often overlapping preceding and distinctly bicarinate, clypeal carinae often succeeding rows (one queen) ...... produced as small denticles, if anterior ...... Cardiocondyla nuda (Mayr) clypeal margin rounded, then eye large .....109

Head and mesosoma of worker moderately 109. Colour brown to blackish in Barrow Island shining, the underlying microreticulate populations; erect and suberect setae very pattern effaced in places, particularly on the rarely present on head and alitrunk; viewed humeral angles, the petiolar node and the in profile, promesonotum flattened and postpetiole; in rear view, appressed gastral truncated (Figure 179) ...... setae shorter, often separated from preceding ...... Monomorium sydneyense Forel and succeeding rows by 0.5-1 x their own Colour yellow or dingy yellow; erect and length ...... Cardiocondyla atalanta Forel suberect setae often present at humeral 104. Antenna with 11 segments ...... 105 angles; viewed in profile, promesonotum often rounded, more elongate (Figure 180) Antenna with 12 segments ...... 112 ...... 110

105. Viewed in profile, eye distinctly oblique, often 110. Eye small, elliptical, eye length ≈ 1.15 reaching to venter of head capsule, distance x greatest width of antennal scape; six from head capsule usually much less than erect setae present on promesonotum and length of eye (Figure 175) ...... propodeum ≥; dingy yellow with brownish- ...... Monomorium eremophilum Heterick yellow gaster ...... Monomorium ‘antipodum’

Viewed in profile, eye situated along Eye larger, tending to elongate, eye length 1.75 longitudinal axis of head, distance from x greatest width of antennal scape ≥; erect mandible at most only slightly less than mesosomal setae absent or present at humeral length of eye (Figure 176) ...... 106 angles only; concolorous yellow ants ...... 111 ANTS OF BARROW ISLAND 399

FIGURE 170 FIGURE 171 FIGURE 172

FIGURE 173 FIGURE 174 FIGURE 175 FIGURE 176

FIGURE 177 FIGURE 178

FIGURE 179 FIGURE 180 400 B.E. HETERICK

111. Erect and suberect setae always present on In profile, indentation of ventral surface of body, usually presenting as a particularly petiole under the node just before its junction prominent pair on the humeral angles and with postpetiole barely discernible; petiolar always as a pair of setae directed posteriad node not obviously arched, its anterior and posterior faces approximately equal in length on the petiolar node and postpetiole, erect (Figure 188) ...... and suberect setae also present on gastral ...... Monomorium rubriceps group sp. JDM 1175 tergites (Figure 181); in some workers small setae that are normally appressed are 115. Head and mesosoma finely microreticulate, subdecumbent or semierect on body and the ant generally weakly shining; eye large, antennal scapes, giving the ant a fuzzy eye length ≈ 3 x greatest width of antennal appearance; propodeum and katepisternum scape ...... of mesopleuron usually with sculpture ...... Monomorium punctulatum Heterick reduced to a few striae ...... Monomorium disetigerum Heterick Head and anterior pronotum smooth and glossy with katepisternum and lower Erect and suberect setae always absent from propodeal flanks reticulate and moderately body (Figure 182); propodeum typically shining; eye rather small, eye length ≈ 1.5 x shagreenate and some shagreenation usually greatest width of antennal scape in Western Australian populations ...... evident on mesopleuron or, at least, on ...... Monomorium leae Forel katepisternum...... Monomorium sydneyense complex sp. JDM 101 116. Larger species (HW of minor worker ≈ 0.8 mm); humeral angles in minor worker 112. Basal tooth much broader than other denoted by a small but distinct denticle preapical teeth (Figure 183); distinctly (major worker unknown) (Figure 189) ...... polymorphic, with large-headed major ...... Pheidole sp. JDM 684 workers having rather small eyes...... Monomorium euryodon Heterick Smaller species (HW of minor worker ≤ 0.7 mm); humeral angles unarmed (Figure 190) Basal tooth of same size or smaller than ...... 117 other preapical teeth (Figure 184); worker monomorphic or exhibiting monophasic 117. Minor worker matt, uniformly allometry ...... 113 microreticulate, and with striae and cross ribs on the head and mesosoma mostly only weakly indicated, if at all; propodeum 113. In profile, postpetiole massively developed weakly longitudinally bicarinate, the carinae and visibly much thicker than petiolar node; separating dorsal and lateral surfaces in anteroventral postpetiolar process a large, minor and major worker; viewed from above, conspicuously protruding lip (Figure 185) occipital lobes of major workers smooth and ...... 114 glossy, without transverse rugae (Figure 191) ...... Pheidole sp. JDM 536 In profile, postpetiole not massive, approximately as thick through as petiolar Head and mesosoma of minor worker node; anteroventral postpetiolar process an variously sculptured (often with smooth, inconspicuous ledge (Figure 186) ...... 115 shining patches) or sculpture completely lacking, but where present, always including some distinct striae and cross ribs; lateral 114. In profile, ventral surface of petiole under carina usually lacking on propodeum or the node conspicuously indented just before incomplete and formed by one or more its junction with postpetiole; petiolar node longitudinal striae; viewed from above arched posteriad, its anterior face decidedly occipital lobes of known major workers longer than its posterior face (Figure 187) ...... always with variably developed transverse ...... Monomorium insolescens Wheeler rugae (Figure 192) ...... 118 ANTS OF BARROW ISLAND 401

FIGURE 181 FIGURE 182 FIGURE 183

FIGURE 184 FIGURE 185 FIGURE 186

FIGURE 187 FIGURE 188 FIGURE 189

FIGURE 190 FIGURE 191 FIGURE 192 402 B.E. HETERICK

FIGURE 193 FIGURE 194

118. Minor workers yellow, the lateral humeral REFERENCES area of the promesonotum smooth and Australian Broadcasting Corporation (2011). (http:// shining; in full-face view, frons almost www.abc.net.au/nature/island/ep3/about1.htm). smooth apart from longitudinal striae, Accessed 22 February 2013. (Used with permission.) shining; in dorsal view, occipital lobes of Bureau of Meteorology (http://www.bom.gov.au/ major worker slightly flattened, with strong climate/averages/tables/cw_005058.shtml). Accessed transverse rugae at the apex of the lobes 22 February 2013. (Figure 193)...... Pheidole sp. JDM 1134 Bamford, M.J. and Bamford, A.R. (2005). Gorgon Minor workers usually shades of brown, but Development on Barrow Island Technical Report – in paler, more tawny specimens, lateral Mammals and Reptiles. Technical Appendix C2 to Draft EIS/ERMP for the Proposed Gorgon Development. humeral area of promesonotum almost Chevron Australia, Perth, Australia. always uniformly microreticulate; frons with Callan, S.K., Majer, J.D. and Moro, D. (2011). microsculpture (such as microreticulation) Documenting the terrestrial invertebrate fauna of as well as longitudinal striae, and mainly Barrow Island. Australian Journal of Entomology 50: matt; in dorsal view, occipital lobes of major 323−343. worker evenly rounded, the apical rugae Clark, J. (1924a). Australian Formicidae. Journal of the often weaker than those lower down, or even Proceedings of the Royal Society of Western Australia 9: absent (Figure 194) ...... 119 72−89. 119. Larger species (minor worker head width Clark, J. (1924b). Australian Formicidae. Journal of the ≈ 0.6 mm); antennal scape longer (exceeds Proceedings of the Royal Society of Western Australia 10: 75−89. vertex by ≥ 1 x its greatest width) (major worker not held in JDM Collection) ...... Clark, J. (1926). Australian Formicidae. Journal of the Royal Society of Western Australia 12: 43−51...... Pheidole mjobergi Forel Clark, J. (1930a). Some new Australian Formicidae. Smaller species (minor worker head width Proceedings of the Royal Society of Victoria (n.s.) 42: ≤ 0.5 mm); antennal scape shorter (exceeds 116−128. vertex by ≤ 0.5 x its greatest width) ...... Clark, J. (1930b). The Australian ants of the ...... Pheidole sp. JDM 177 (nr variabilis Mayr) Dolichoderus (Formicidae). Subgenus Hypoclinea Mayr. Australian Zoologist 6: 252−268. Clark, J. (1934). Notes on Australian ants, with ACKNOWLEDGEMENTS descriptions of new species and a new genus. Memoirs of the National Museum of Victoria 8: 5−20. I wish to thank Dr Nihara Gunawardene Clark, J. (1936). A revision of Australian species of for the kind sharing of her Barrow Island Ant Rhytidoponera Mayr (Formicidae). Memoirs of the database, for testing the ant taxonomic key National Museum of Victoria 9: 14−89. and for statistical advice. Thanks also go to Dr Clark, J. (1938). The Sir Joseph Banks Islands. Reports Christopher Taylor, who also tested the taxonomic of the McCoy Society for Field Investigation and key. The use of many Figures produced by Mr Research. Part 10. Formicidae (Hymenoptera). Brad Durrant and previously appearing in Records Proceedings of the Royal Society of Victoria (n. s.) 50: of the Western Australian Museum Supplement No. 356−382. 76 is acknowledged. Professor Jonathan Majer Clark, J. (1943). A revision of the genus Promyrmecia commented on an initial draft. I also wish to thank Emery (Formicidae). Memoirs of the National Museum of the contributions of the two referees (Dr Steve Victoria 13: 83−149. Shattuck and Dr Phil Ward) who commented on the Clark, J. (1951). The Formicidae of Australia. Vol. 1. manuscript when it was submitted. Subfamily Myrmeciinae. CSIRO, Melbourne. 230 pp. ANTS OF BARROW ISLAND 403

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A new species of the genus 137−163. Meranoplus. Memoirs of the Queensland Museum 13: 148−150. MANUSCRIPT RECEIVED 8 AUGUST 2012; ACCEPTED 20 MARCH 2013. 404 B.E. HETERICK ADDENDUM Since this paper was compiled and submitted for review, two additional ant species have been discovered on the Island. Iridomyrmex bicknelli Emery is a gracile, iridescent species that is common in much of southern Australia, but is much rarer north of the Tropic of Capricorn. The ant will come out in the taxonomic key at couplet 65, and can easily be distinguished from I. anceps and I. minor by its uniform dark grey colouration and bluish or yellowish-green iridescence. Melophorus sp. JDM 951 will come out at couplet 41, where it can best be separated from M. turneri in respect of its major caste, which is distinguished by stout, incurved mandibles that are likely used for milling seed (the mandibles not noticeably stout or incurved in M. turneri major workers). The minor workers of Melophorus sp. JDM 951 are matt and dark brown in colour (the cuticle normally lighter in colour and glossier in M. turneri) and the appressed setae on the gaster are relatively long and overlapping (short and well-separated in M. turneri). This species is uncommon and appears to have a localised distribution in northern Australia. Shortly before the proofs came out, the revision of Polyrhachis (subgenus Hagiomyrma) by Kohout was published. There is some doubt about the identity of ‘melanura’, as the Barrow Island specimens do not quite fit the description in the published key, and occur considerably further south than the material examined by Kohout.