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C S I R O P U B L I S H I N G

Australian Journal of Zoology

Volume 45, 1997© CSIRO Australia 1997

A journal for the publication of the results of original scientific research in all branches of zoology,except the taxonomy of invertebrates

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A Morphological and Molecular Reviewof Camponotus terebrans (Lowne)(Hymenoptera : Formicidae)

A. J. McArthurA, M. AdamsA and S. O. ShattuckB

ASouth Australian Museum, North Terrace, Adelaide, SA 5000, Australia.BCSIRO Entomology, PO Box 1700, Canberra, ACT 2601, Australia.

Abstract

Camponotus terebrans (Lowne) of the subfamily Formicinae is reviewed with the aid of both morphologicaland allozyme techniques. C. testaceipes (Smith) and C. darlingtoni Wheeler are previously recognisedsynonyms while C. latrunculus victoriensis Santschi and C. myoporus Clark are newly proposed synonyms.C. darlingtoni is removed from synonymy and its lectotype is designated. Although considerable variationhas been recognised in this species, this variation is considered to be clinal. A description of this variablespecies and its known distribution is presented. Examples of its habitat preferences and behaviour are alsoincluded.

Introduction

There are more than 100 described species of Camponotus in Australia, yet few can beidentified with certainty. Most species can be identified only by comparing specimens with typematerial. This is necessary because of the paucity and ambiguity of original descriptions and thelack of a published key for the genus.

Camponotus terebrans (Lowne) is one of the most common ants in sandy soils of southernAustralia and is one of the first ant species to colonise disturbed sites. Its castes display widevariation in size, colour, pilosity and profile, and because of this variation, major and minorworkers may conceivably be misidentified as different species if collected away from the nest.Over the last decade or so, ecologists have recognised that the number of ant taxa present at asite is an indicator of ‘environmental success’, particularly when a disturbed site is beingrevegetated (Majer 1983). In monitoring these sites, field biologists have been frustrated by alack of taxonomic literature. The aims of this paper are to clarify the taxonomy of this variablespecies by a combination of morphological and molecular analysis, to present a cleardescription, and to provide an overview of the biology of the species.

Materials and MethodsAbbreviations

Location of material examined

ANIC Australian National Insect Collection, Canberra, Australian Capital TerritoryBMNH Natural History Museum, London, UK

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USANMBA Naturhistorisches Museum, Basel, SwitzerlandNVMA Museum of Victoria, Melbourne, VictoriaSAMA South Australian Museum, Adelaide, South AustraliaWAM Western Australian Museum, Perth, Western Australia

Australian Journal of Zoology, 1997, 45, 579–598

10.1071/ZO97055 0004-959X/97/060579$05.00

© CSIRO Australia 1997

Morphological Analysis

Many morphological characters were examined in an attempt to find those most suitable for taxonomicanalysis. When conducting microscopic examination, pilosity was best observed by transmitted light (lightsource below the specimen), whereas reflected light was better for other observations. For each workercaste, characters that scored the greatest apparent variation were as follows: (1) eye length relative to headwidth, (2) mid-tibial length relative to head width, (3) colour of head, (4) colour of mesosoma, (5) propodealpubescence, (6) number and length of setae on propodeal dorsum, and (7) curvature of the propodeum andmetanotum. Voucher specimens representing all nests examined in this study have been deposited with theSouth Australian Museum.

Allozyme Analyses

The protocols used here follow McArthur and Adams (1996). Briefly, animals were placed within‘Eppendorf’ tubes, snap-frozen in the field with liquid nitrogen, and stored at –70°C until required. Once allsamples had been collected, each ant was thawed, examined under a dissecting microscope to confirm itsidentity, and the head and mesosoma homogenised by hand before being loaded onto cellulose–acetate gels(‘Cellogel’, Chemetron, Milan). A total of 43 ants from 25 nests was run in an initial screen of all enzymesfound to be suitable by McArthur and Adams (1996). Following this initial screen, nests from additionalregions in South Australia were sampled in order to further explore the patterns of genetic andmorphological variation observed. These additional ants were analysed only for the most informativegenetic markers.

Allozyme electrophoresis was carried out as detailed by Richardson et al. (1986). The followingenzymes were successfully screened: aconitate hydratase (ACON, EC 4.2.1.3), aminoacylase (ACYC, EC3.5.1.14), fructose-bisphosphate aldolase (ALD, EC 4.1.2.13), arginine kinase (ARGK, EC 2.7.3.3), enolase(ENOL, EC 4.2.1.11), esterase (EST, EC 3.1.1.), fructose-bisphosphatase (FDP, EC 3.1.3.11), fumaratehydratase (FUM, EC 4.2.1.2), glyceraldehyde-3-phosphate dehydrogenase (GAPD, EC 1.2.1.12), aspartateaminotransferase (GOT, EC 2.6.1.1), glucose-6-phosphate isomerase (GPI, EC 5.3.1.9), alanineaminotransferase (GPT, EC 2.6.1.2), hexokinase (HK, EC 2.7.1.1), isocitrate dehydrogenase (IDH, EC1.1.1.42), L-lactate dehydrogenase (LDH, EC 1.1.1.27), ‘malic’ enzyme (ME, EC 1.1.1.40), nucleoside-diphosphate kinase (NDPK, EC 2.7.4.6), dipeptidase (PEPA, EC 3.4.13.), tripeptide aminopeptidase (PEPB,EC 3.4.11.), proline dipeptidase (PEPD, EC 3.4.13.), phosphoglycerate kinase (PGK, EC 2.7.2.3),phosphoglucomutase (PGM, EC 5.4.2.2), pyruvate kinase (PK, EC 2.7.1.40), superoxide dismutase (SOD,EC 1.15.1.1) and triose-phosphate isomerase (TPI, EC 5.3.1.1). The conventions for referring to loci andallozymes follow Adams et al. (1987).

580 A. J. McArthur et al.

Collectors of material examined

AAT A. AtkinsAHB A. H. BurbidgeAMA A. J. McArthurALY A. L. YenANA A. N. AndersenANZ Australia and New Zealand Scientific

Exploration Society Inc.BBL B. B. Lowery

CHW C. H. WattsDH D. Hirst

EGM E. G. MathewsGFG G. F. GrossGLH G. L. HowieJAF J. A. ForrestJAH J. A. Herridge

JC J. ClarkJDE J. D. ErskineJEF J. E. Feehan

LHU L. HuntMAA M. AdamsMLS M. L. Simpson

NBT N. B. TindalePJF P. J. Fargher

PJMG P. J. M. GreensladeRGR R. GrundRHF R. H. Fisher

RH R. HayRPF R. P. Field

RWT R. W. TaylorSANPNS South Australian National Parks and

Wildlife Nullarbor SurveySANPSOPS South Australian National Parks and

Wildlife South Olary Plains SurveySANPSDS South Australian National Parks and

Wildlife Sturt Stony Desert SurveySANPVS South Australian National Parks and

Wildlife Vertebrate SurveySEG Scientific Expedition Group

TG T. GreavesTHE T. Herbert

TR T. ReardonWMA W. M. McArthur

The genetic-divergence values between nests were initially calculated as percent fixed differences(%FD) (see Richardson et al. 1986) and depicted visually by a dendrogram constructed with the unweightedpair group method of analysis (UPGMA) (Sneath and Sokal 1973). Subsequently, the multivariate techniqueof principal co-ordinates analysis (PCoA), as implemented by the statistical package PATN (PatternAnalysis Package) (Belbin 1987), was used to display the genetic relationships amongst all South Australiannests examined, using Rogers’ genetic distances (Rogers’ R) (Rogers 1972) to construct the input matrix.The rationale for this approach is explained by McArthur and Adams (1996).

Results

Morphological Analysis

Several morphological characters show considerable variation. These include colour, theprofiles of the propodeum and metanotum, and the lengths of the eyes and mid-tibiae (Fig. 1).Details of these characters are as follows.

The colour of the mesonotum varies from dark in southern and western regions to light ininland areas. This change in colour is gradual and uniform between the extremes of the range,and light and dark individuals have never been found in sympatry. The size of the eyes (Fig. 2)and the mid-tibia (Fig. 3) show a distinct north–south clinal pattern similar to that found in bodycolour. In southern regions the eyes are relatively smaller while in northern areas the eyes arelarger and legs are longer. The variation seen in these characters shows gradual and continualchange across these areas and is considered clinal.

The variation in the curvature of the metanotum and propodeum is most noticeable in majorworkers. This surface varies from straight to feebly convex and intra-nest variation can begreater than inter-nest variation. In all major workers the metanotum itself is distinct but variable(particularly when viewed from the side). The metanotum fades as size decreases and iscompletely absent in minor workers. No geographic patterns were detected in the variation ofthese characters.

581Review of Camponotus terebrans

Fig. 1. Medium worker of Camponotus terebrans indicating characters that are referred to in the text. Theupper portion is in lateral view. The leg is drawn rotated 90° at the coxa–femur joint and is viewed from thefront to show the coarse bristles on the inside surface of the tibia. Scale line, 1 mm.


Fig. 2. Relationship between eye length and head width for three localities along a north–southtraverse.

Fig. 3. Relationship between mid-tibia length and head width for four localities along a north–southtraverse.

Systematic Implications of the Allozyme Data

The allozyme profiles at 32 loci for the 43 ants, representing 25 nests, included in the initialscreen are shown in Table 1; the geographic localities of these nests are shown in Fig. 4. Sixteenloci displayed allozyme variation, with a maximum of four allozymes found at each of severalloci. Whilst within-nest polymorphism was common, all 68 cases involved the presence of onlytwo allozymes, and in only one of these cases were no heterozygotes detected (Acon-1 for Nest23). As such, it is appropriate to employ our previous approach of using the nest as the unit ofanalysis (McArthur and Adams 1996).

The genetic relationships amongst the 25 nests, based on the pairwise comparison of nests for%FDs, are displayed in Fig. 5. Overall, nests show a high degree of genetic similarity across thegeographic range sampled, with a maximum of 13%FD found between any two nests, and anaverage level of divergence of 5.4%FD (the genetic-distance matrix is not shown). Nevertheless,two major groupings are evident in Fig. 5, differing at an average of 9.1%FD (equating toapproximately three fixed differences). Most significantly, the geographic distribution of the twogroups is distinctly non-random, and reflects a simple, underlying geographic pattern. First,there is a ‘southern’ group (Nests 1–8) consisting of all of the Western Australian nests exceptthe two from Kalgoorlie, plus those from Fleurieu Peninsula, South Australia (Nests 21–25). Thesecond group consists of the ‘northern’ nests, namely the two from Kalgoorlie (Nests 9, 10) plusthe remaining six northern locations from South Australia (Nests 11–20). An examination ofTable 1 indicates that the northern and southern groups can be diagnosed unequivocally by theirallozyme profiles at the loci Est-2 and Acon-1 (although only Est-2 is fully diagnostic on itsown), with further differentiation within either Western Australia or South Australia apparent atthe loci Enol and Est-1.

No other major genetic discontinuities are evident from the allozyme data, with only a singlelocus per group showing any evidence of regional differentiation within each of the southern andnorthern groups. For the southern group a comparison of the Western Australian with the SouthAustralian nests reveals a near-fixed difference at the locus Enol (Table 1). The northern groupalso shows within-group divergence at one locus, with the Kalgoorlie nests possessing a near-fixed difference at Est-1 from those in South Australia (Table 1). The genetic divergence at thesetwo loci is responsible for the primary dichotomies found within each group, as displayed in Fig.6. Thus, the genetic divergence within each group across a geographic range of more than 1000km is clearly less than that displayed between the northern and southern groups, which weresampled to within 200 km of one another (e.g. Esperance and Kalgoorlie in Western Australia,Cox Scrub and Calperum in South Australia).

In an attempt to resolve the ambiguous pattern of genetic diversity observed in C. terebrans,additional nests were sampled throughout eastern South Australia from the Riverland regionsouthwards. All available blocks of sandy habitat were visited in an effort to locate additionalnests. In this manner, nine nests from a further seven sites were sampled, with these additionalants being analysed only for the enzymes ACON, EST, ENOL, GPI, PEPB and TPI, the mostinformative genetic markers from the initial screen. The results of this screen are presented inTable 2.

Even though the formerly diagnostic locus Est-1 now displays a few heterozygotes, onceagain the overall allozyme profiles at Acon-1, Est-1 and Est-2 reveal a distinction between thetwo northern-type nests (Nests 33 and 34 from Stockyard Plain) and all others to the south,including Nest 32 from Billiat Conservation Park, the first patch of sandy terrain to the south.The maintenance of the dichotomy between the northern and southern groups is betterdemonstrated by a PCoA analysis on the Rogers’ genetic-distance matrix of pairwisecomparisons of all of the 24 nests sampled in South Australia (data from the nine loci run for allnests and extracted from Tables 1 and 2). The PCoA scores for the first two dimensionsextracted in such an analysis are plotted in Fig. 6. The distinction between the northern andsouthern groups is clearly shown, with all nests falling within either one or the other of the twoclusters. Thus there is reasonable genetic integrity within each of the two groups across


584A

. J. McA

rthur et al.

Table 1. Allozyme profiles of the 43 ants examined in the initial screenAnts are designated by nest Nos 1–25 (localities as per Fig. 4) and also alphabetically, where two or more ants from the nest were sampled. A dashindicates no activity for that particular enzyme. Loci: 1, Acon-1; 2, Acon-2; 3, Acyc; 4, Ald; 5, Argk; 6, Enol; 7, Est-1; 8, Est-2; 9, Fdp-1; 10, Fdp-2; 11,Fum; 12, Gapd; 13, Got-1; 14, Got-2; 15, Gpi; 16, Gpt; 17, Hk; 18, Idh; 19, Ldh; 20, Me; 21, Ndpk; 22, PepA; 23, PepB-1; 24, PepB-2; 25, PepD-1; 26,

PepD-2; 27, Pgk; 28, Pgm-1; 29, Pgm-2; 30, Pk; 31, Sod; 32, Tpi

Ant Locus

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32

1a c b a a a a a a a a a a a a bc a a a b c b a a a a cd a a a b a b 1b c b a - a a - a a - - a a - - - a a - ac b - a a a cd a a - b a b 2a c b a a a a a a a a a a a a b a a a b c b a a a a cd a a a b a b 2b c b a a a a a a a a a a a a b a a a b c b a a a a d a a a b a b 3a c b ab a a a a a a a a a a a b a a a b c ab a ab a a cd a a a b a b 3b c b ab - a a a a a a a a a a - - a a - c ab - b a a d a a a ab a b 4a - - ab a - a - a a - - - - - - a a a - c b - a - a d - a a b a -4b c b a a a ab - a a a a a a a b a a a b c b a ab a a bd a a a b a b 4c c b a a a a b a a a a a a a bc a a a b c b a ab a a d a a a b a b 5a ac b a a a a a a a a a a a a b a a a b c b a b a a d a a a b a b 5b ac b a a a a ab a a a a a a a b a a a b c b a bc a a d a a a b a b 6a c b a a a a a a a a a a a a b a a a b c b a b a a d a a a b a b 6b c b a a a a a a a a a a a a b a a a b c b a ab a a d a a a b a b 7 cd b a - a a a a a a a a a a ab a a a b c b a ab a a d a a a b a b 8a c ab a a a a a a a a a a a a ab a a a b c b ab ab a a cd a a a b a bd 8b c b a a a a ab a a a a a a a ab a a a b c b - ab a a cd a a a b a b 8c c ab a a a a a a a a a a a a ab a a a b c b ab ab a a cd a a a b a bd 9a - - a - a b a b a a a a a a b a a a b c b a a - a ad a a a b a b 9b d b a a a b a b a a a a a a b a a a b c b a a - a d a a a b a b

585R

eview of C

amponotus terebrans

10a d b a a a b a b a a a a a a b a a a b c b a a a a ad a a a b a b 10b d b a a a b a b a a a a a a b a a a b c bc a a a a ad a a a b a b 11a d b a a a b b bc a a a a a a b a a a b c b ab ab a a d a ab a b a b 11b d b a a a b b b a a a a a a b a a a ab ac b a a a a d a a a b a b 12 d b a a a b ab b a a a a a a b a a a b c b a a a a d a a a b a b 13 d b a a a b b b a a a a a a b a a a b c b a a a a d a ab a b a b 14 d b a a a b b b a a a a a a b a a a b c b a a a a d a ab a b a b 15a d b a a a b b b a a a a a a b a a a ab c b a a - a d a a a b a b 15b cd b a a a b b b a a a a a a b a a a a c b a ab a a d a a a b a b 16a d b a a a b ab b a a a a a a b a a a b c b a ab a a d a a a b a b 16b - - - - - b b b a a - a - - b - a - - c b a a - a d - a - b a b 17a d b a - a b b b a a a a a a b a a - b cd b a a a a d a a a b a b 17b d b - a a b b b a a a a a a b a a a b cd b a a a a d a a a b a b 18a - b - - - b - b a a - a - a - - a a - bc b - b a a d - a - b a -18b cd b a a a b b bc a a a a a a b a a a b c b a b a a d a a - b a b 18c d b a a a b b bc a a a a a a b a a a b bc b a b a a d a ab a b a b 19 d b a a a b b b a a a a a a b a a a b c b a ab a a d a a a bc a bc 20 d b a a a b - bc a a a a a a b a a a - cd b a ab a a d a a - bc a b 21 bc b a a a b a a a a a a a a bd a a a b c b a b a a d a a a b a b 22a bc b a a a b a a a a a a a a bd a a a b c b a ab a a d a a a b a b 22b b b a a a b a a a a a a a a d a a a b c b a b a a d a a a b a b 23a b b a a a b a a a a a a a a d a a a b c b a b a a d a a a b a b 23b c b a a a b a a a a a a a a bd a a a b c b a ab a a d a a a b a b 24 bc b a a a b a a a a a a a a b a a a b c b a ab a a d a a a b a b 25 c b a a a b a a a a a a a a b a a a b c b a b a a d a a a b a ab

distances of roughly 300 km in the case of the southern group (e.g. Billiat to Beachport) and600 km for the northern nests (e.g. Mildura to Lake Eyre), albeit based largely on the results ofthree loci, none of which is fully diagnostic when applied individually.

Integration of the Morphological and Allozyme Analyses

The allozyme data suggest the presence of two genetic groups within C. terebrans, broadlycorresponding to a northern and a southern group. The two groups are genetically similar, withlevels of divergence between adjacent regions of 3–6%FD, based mainly on combinations ofnearly fixed differences at three loci (Table 1). Our attempts to find these two forms in sympatryin South Australia were not successful, although they do occur as close as 100 km on adjacentblocks of sandy terrain (Billiat and Stockyard Plain). However, as only a relatively small numberof nests was sampled, it may be that more-robust estimates of allele frequency for populationswould reveal the presence of all key northern and southern alleles in this geographicallyintermediate region. We were unable to conduct detailed sampling in Western Australia, where atransect from Esperance to Kalgoorlie would be highly desirable as it would include both forms.


Fig. 4. Specimens for the allozyme study were collected and stored in liquid nitrogen atlocalities indicated on the map. Numbers on the map refer to the localities named below.Numbers in parentheses refer to columns headed ‘Ant’ in Tables 1 and 2. Aldinga, 21(21–24); Beachport, 26 (26); Billiatt, 32 (32); Calperum, 20 (20, 21); Cox Scrub, 24 (24, 25);Denmark, 5 (5); Esperance, 8 (8, 9); Kalgoorlie, 10 (10, 11); Koortanyaninna Creek, 16 (16,17); Lake Eyre South, 12 (12); Marree, 13 (13–15); Mildura, 18 (18); Mount Rescue, 29(29); Mount Hope, 27 (27); Ngarkat, 30 (30, 31); North Bannister, 1 (1–3); Porongurup, 6 (6,7); Stockyard Plain, 33 (33, 34); Walpole, 4 (4); West Avenue Range, 28 (28); YaningurieWaterhole, 19 (19).

This lack of sympatric associations of the genetic types is consistent with morphology, wherebody colour and eye size show a similar pattern.

While the level of genetic and morphological divergence found between the northern andsouthern groups of these ants suggests that they may represent distinct taxa, it is not of sufficientmagnitude in allopatry to draw the definite conclusion that they represent distinct biologicalspecies. For example, it is possible that the two groups represent the extremes of a north–southcline, or that they are formerly disjunct subspecies that may be in the process of merging into asingle metapopulation. Further sampling in geographically intermediate regions is needed toresolve this borderline systematic scenario.

Systematics

Camponotus terebrans (Lowne)

Formica testaceipes Smith, 1858: 39 (preoccupied by Leach, 1825: 290).Camponotus testaceipes. – Mayr, 1861: 60 (new comb.).Formica terebrans Lowne, 1865: 278.Camponotus (Myrmophyma) testaceipes. – Forel, 1914: 269.Camponotus testaceipes. – Emery, 1925: 102.Camponotus latrunculus victoriensis Santschi, 1928: 479 (new syn.).Camponotus myoporus Clark, 1938: 379 (new syn.).

Material Examined

Types examined

Formica terebrans. BMNH, Austr Lowne, F. Smith coll 79/22. Two minor workers on one card.HW = 1.7 mm, HL = 1.9 mm, EL = 0.5 mm. HW = 1.4 mm, HL = 1.7 mm, EL = 0.4 mm. Head andmesosoma brownish, legs yellowish. One medium worker on card.


Fig. 5. Dendrogram depicting the genetic relationships among the 25 Camponotus terebrans nestssampled in the initial allozyme analysis. SA, South Australia; WA, Western Australia.

Formica testaceipes. BMNH Drawer 11-486, F. testaceipes Type/ Sm. BM Type 11-619. K Geo S. 5615. One major worker pinned: HW = 3.3 mm, HL = 3.0 mm, EL = 0.6 mm. Head and mesosoma brown,legs yellow.

Camponotus latrunculus victoriensis. NMBA Type Victoria Elsternwick Barrett Sammlung Dr F.Santschi Kairouan. One major, 1 minor worker and 1 male on one pin. HW = 3.0 mm, HL = 2.8 mm, EL =0.67 mm.


Fig. 6. Plot of the principal co-ordinates scores for the first two dimensions of a Principal Co-ordinatesAnalysis (PCoA) for all 24 nests sampled from South Australia and Victoria. Localities of nests are asfollows: 11, Kalgoorlie; 12, Lake Eyre South; 13–15, Marree; 16, 17, Koortanyaninna Creek; 18, Mildura;19, Yaningurie Waterhole; 20, Calperum; 21–24, Aldinga; 25, Cox Scrub; 26, Beachport; 27, Mount Hope;28, West Avenue Range; 29, Mount Rescue; 30, 31, Ngarkat; 32, Billiatt; 33, 34, Stockyard Plain. The firsttwo dimensions of the PCoA account for 71% of the total multivariate variation present in the originalmatrix.

Camponotus latrunculus victoriensis. NMBA Type Victoria Belgrave Barrett Sammlung Dr. FSantschi Kairouan. One major, 1 medium and 1 minor on 1 card. HW = 2.5 mm, HL = 2.45 mm, EL = 0.58mm. HW = 1.3 mm, HL = 1.6 mm, EL = 0.44 mm.

Camponotus myoporus. NMVA Reevesby I. J. Clark. 1 major worker, 1 minor worker. HW = 2.8 mm,HL = 2.7 mm, HW = 1.9 mm, HL = 2.1 mm.

Type specimens of C. latrunculus victoriensis and C. myoporus examined during this study conformwith the diagnosis of C. terebrans proposed here, thus establishing their synonymy with this species.

Other material examined

Northern Territory: New Crown HS, 1992, JEF (SAMA). New South Wales: Botany Bay, H. P.Schrader (ANIC); Fort Grey, Sturt Natl Pk, 1995, DH (SAMA); Lake Cawndilla, 1971, BBL (ANIC); MyallLakes, 1977, M. Fox (ANIC); Wilcannia, 47 mi E, 1947, TG (ANIC); Yanco, W. W. Froggatt (SAMA).Queensland: Windorah, 50 km W, 1996, Anon. (SAMA); Annandale, 125 km NW Birdsville, 1993, A. W.


Table 2 Allozyme profiles of an additional 17 ants from seven nests in South Australia, south of theRiverland

The profiles of four ants taken from nests already screened (Nests 21 and 24, southern group, and Nest 11,northern group) are also presented. Ants are designated by nest Nos 11–34 (localities as per Fig. 4) and alsoalphabetically, where two or more ants from the nest were sampled. A dash indicates no activity for thatparticular enzyme. Loci: 1, Acon-1; 2, Acon-2; 6, Enol; 7, Est-1; 8, Est-2; 15, Gpi; 23, PepB-1; 24, PepB-2;

32, Tpi

Ant Locality Locus

1 2 6 7 8 15 23 24 32

21b Aldinga c b b a a b b a b24b Cox Scrub c b b a a b a a ab24c Cox Scrub c b b a a bd b a b26a Beachport c b b a a b b a ab26b Beachport c b b a a b b a ab27a Mt Hope c b ab a b a b a ab27b Mt Hope c b b a a b b a a28a West Avenue Range c b b a a b b a ab28b West Avenue Range c b b a a bd b a ab29a Mt Rescue c b b a a b ab a b29b Mt Rescue c b b a a bd – a b30 Ngarkat c b b – a b b a b26b Beachport c b b a a b b a ab27a Mt Hope c b ab a b a b a ab27b Mt Hope c b b a a b b a a28a West Avenue Range c b b a a b b a ab28b West Avenue Range c b b a a bd b a ab29a Mt Rescue c b b a a b ab a b29b Mt Rescue c b b a a bd – a b30 Ngarkat c b b – a b b a b31 Ngarkat c b b a a b b a b32a Billiat c b b a a b b a b32b Billiat c b b a a b b a b32c Billiat c b b a a b b a b33a Stockyard Plain d b b ab b b b a b33b Stockyard Plain – b b ab b b b a b34a Stockyard Plain – b b ab ab b b a b34b Stockyard Plain d b b b b b b a b11c Lake Eyre South d b b ab ab b ab a b

Lewis (SAMA); Diamantina Ridge, A. M. Morgan (SAMA); Birdsville, 20 mi WNW, 1973, C. W.McCubbin (SAMA). South Australia: Abyssinia Bay, Canunda Cons. Pk, 1996, AMA (SAMA);Aldinga Beach, 1969, BBL (ANIC); Aldinga Scrub, 1987, DH (SAMA); Aldinga Sellicks Beach Res., 1987,EGM and JAF (SAMA); Aldinga Scrub, 1995, AMA (SAMA); Aldinga Scrub, with Ogyris, 1995, RGR(SAMA); Andamooka Range, 1947, GFG and J. H. Mitchell (SAMA); Anna Ck, 37 km W, 1975, EGM(SAMA); Arrabury, 10 km SW, 1975, PJMG (ANIC); Banff, Coorong Cons. Pk, 1975, PJMG (ANIC);Beachport, 1980, L. Queale (SAMA); Beachport Two Lakes, 1988, AMA (SAMA); Billiat Cons. Pk, 2 kmS, 1996, AMA (SAMA); Calperum, with Ogyris, 1995, S. Xuereb (NMVA); Calperum, Amalia, 1995,AMA (SAMA); Calperum HS, 1995, AMA (SAMA); Calperum, Murphys, 1995, AMA (SAMA);Calperum, NE corner, 1995, AMA (SAMA); Canunda Cons. Pk, nr beach, 1988, AMA (SAMA); CanundaCons. Pk, South End, 1988, AMA (SAMA); Canunda Cons. Pk, 1992, AMA (SAMA); Carribie York Pen.,NBT (SAMA); Ceduna, 1957, BBL (ANIC); Ceduna, 20 km E, 1988, JAF (SAMA); Clayton Crossing,1955, E. T. Giles (SAMA); Clayton Ridge, 1972, PJMG (SAMA); Coolatoo, Coorong Cons. Pk, 1975,PJMG (ANIC); Cooltong, 1991, AMA and MAA (SAMA); Coonalpyn Byree, 4 km W, 1991, JAF(SAMA); Coongie Lakes, 4.5 km WNW Coongie, 1987, J. Reid (SAMA); Cordillo Downs Invasion Tank,1994, R. Brandle (SAMA); Cortina Stn, 1968, A. W. Forbes (SAMA); Cox Scrub Cons. Pk, 1995, AMA(SAMA); Cullyamurra Waterhole, Coopers Ck, 1990, ANZ (SAMA); Danggali Cons. Pk, 1988, L.Schofield (SAMA); Danggali Cons. Pk, 10 km N Red Tank, 1992, AMA (SAMA); Danggali, Morganvale,1992, AMA (SAMA); Danggali, Morganvale, 1993, R. Ramsay (SAMA); Danggali, NE corner, 1993, AMA(SAMA); Etadunna, Birdsville Track, 12 km SW, 1975, PJMG (ANIC); Ethyl Beach, Yorke Pen., 1990,AMA (SAMA); Etrick Cons. Pk, 1993, GLH (SAMA); Farina, 1916, SAMA Expedition (SAMA); Farina,1994, SANPSDS (SAMA); Flinders I., 1929, F. Wood Jones (SAMA); Flinders I., GFG (SAMA); FurnerDrain M, 1987, AMA (SAMA); Furner Kandara, 1989, AMA (SAMA); Furner Cons. Pk, 1996, AMA(SAMA); Goyder Lagoon, 1957, B. Daly (SAMA); Gum Lagoon, 1992, EGM and JAF (SAMA); HamiltonDowns Stn, 1991, ANZ (SAMA); Hamilton HS dune, 1991, ANZ (SAMA); Innaminka HS, 1957, SAMAExp. (SAMA); Innes Cons. Pk, 1991, RHF (SAMA); Innes Cons. Pk, with Ogyris, 1993, RHF (SAMA);Innes Cons. Pk, 1972, PJMG (ANIC); Innes Cons. Pk, with Ogyris, 1990, RHF (SAMA); Jacobs Range,1988, AMA (SAMA); Johnsons No.3 Bore, 1971, B. K. Head (SAMA); Kangaroo I. (KI), Bayeria Cons. Pk,1994, SANPVS (SAMA); KI, Bayeria Cons. Pk, with Ogyris, 1990, LHU (SAMA); KI, Cape GantheaumeCons. Pk, 1987, GFG and DH (SAMA); KI, Castle Hill Cons. Pk, 1993, AMA (SAMA); KI, Douglas Hill,12 km SSE Karalla, 1990, SANPVS (SAMA); KI, Dudley Cons. Pk, 1990, GFG and DH (SAMA); KI,Dudley Cons. Pk, 1987, SANPVS (SAMA); KI, Dudley Peninsula, 1971, JAF (ANIC); KI, Flinders Chase,1972, PJMG (ANIC); KI, Flinders Chase, Cape du Couedic, 1.6 km NE, 1990, SANPVS (SAMA); KI,Flinders Chase, N boundary, 1992, B. Overton and P. King (SAMA); KI, Flinders Chase, Ravine desCasoars, 6 km N, 1990, EGM and JAF (SAMA); KI, Kelly Hill Caves, 1987, GFG and DH (SAMA); KI,Kelly Hill Cons. Pk, 1990, SANPVS (SAMA); KI, Kiawarra, 1995, THE (SAMA); KI, Kingscote, 1992,SANPVS (SAMA); KI, Kingscote, 9 km S, 1992, SANPVS (SAMA); KI, Mt Thisby, 1990, SANPVS(SAMA); KI, Penneshaw, 13.6 km W, 1990, SANPVS (SAMA); KI, Rocky Ridge, 1972, EGM and JAF(ANIC); KI, Rocky Ridge, 1972, PJMG (ANIC); KI, Salt Lagoon, 1990, SANPVS (SAMA); KI, Seal Bay,1995, THU (SAMA); KI, Seal Bay, 6.5 km N, 1994, THE (SAMA); KI, Snellings Beach, 1993, AMA(SAMA); KI, Vivonne Bay, 1926, SAMA Exp. (SAMA); KI, Western Cove, Nepean Bay, 1987, GFG(SAMA); Kielara Stn, 1990, DH (SAMA); Koortanyaninna Ck, 1994, AMA and JDE (SAMA); KununkaLake, 1970, D. Lacis (SAMA); Lake Eyre South, 1995, MAA (SAMA); Lake Hart, 1994, JAF (SAMA);Lake Meramangye, Victoria Desert, 1976, PJMG (ANIC); Lucindale, Feuerheerdt (SAMA); Madigan Gulf,1956, GFG (SAMA); Marble Range, 1979, GFG (SAMA); Marree, 20 mi S, 1995, MAA (SAMA);McLaren Flat, 1969, C. A. Kirkby (ANIC); Merty Merty, 1993, TR (SAMA); Messent Cons. Pk, 1994,GLH (SAMA); Messent Cons. Pk, with Ogyris, 1996, LHU (SAMA); Mt Hope, 1988, AMA (SAMA);Monteoolina Bore, 1990, ANZ (SAMA); Mt Flint, 1991, P. and I. Gee (SAMA); Mt Rescue Cons. Pk, 1996,AMA and JDE (SAMA); Mt Monster, 1989, AMA (SAMA); Mt Rescue Cons. Pk, Jimmys Well, 1992,EGM and JAF (SAMA); Mungeranie, 1977, JAF (SAMA); Mungeranie, 100 km S, 1993, JAF (SAMA);Murbko, 1973, GFG (SAMA); Myponga, 1969, BBL (ANIC); Myponga, Nixon Skinner Natl Pk, 1969,BBL (SAMA); Ngarkat Cons. Pk, S boundary, 1996, AMA and JDE (SAMA); Ngarkat Cons. Pk, 1993,AMA (SAMA); Nunns Bore, 1990, P. and I. Gee (SAMA); Olympic Dam, 1988, A. Smith (SAMA);Olympic Dam, 1987, EGM and CHW (SAMA); Palankarinna, 1986, J. Thurmer (SAMA); Penola Forest,1995, THE (SAMA); Pertendi, 1991, SANPVS (SAMA); Piccaninnie Ponds, 1982, JAF (SAMA);Pooginook, 1992,AMA and MAA (SAMA); Pooginook, 1992, GLH (SAMA); Port Lincoln, A. M. Lea(SAMA); Port Lincoln, 1960, M. Kenny (SAMA); Port Lincoln, NBT (SAMA); Port Lincoln, SpaldineCove, 1973, PJMG (ANIC); Reevesby I., 1936, D. J. Mahony (ANIC); Reevesby I., 1995, MLS (SAMA);


Renmark, 1980, G. Browning (SAMA); Robe, 1978, PJMG (SAMA); Roneton, 12 km NNE, 1992,SANPSOPS (SAMA); Rotten Swamp, 65 km E Lake Frome, 1977, PJMG (ANIC); Salt Ck, 1994, SAAnimal and Plant Control Commission (SAMA); Salt Ck, 1963, C. T. James (SAMA); Salt Ck, CoorongNatl Pk, 1975, PJMG (ANIC); Scott Cons. Pk, 1996, RGR (SAMA); Scrubby Peak, 1988, AMA and PJF(SAMA); Serpentine Lake, Great Victorian Desert, 1979, PJMG (ANIC); Stirling Range, 1995, PJF(SAMA); Stockyard Plain, 1996, GLH (SAMA); Strzelecki Ck, 1989, AMA (SAMA); Stuart Ck HS, 1975,EGM (SAMA); Sutherlands, 1990, AMA (SAMA); Talia Caves, 1973, JAH (SAMA); Tantanoola,Carpenter Rocks Rd, 1986, AMA (SAMA); Tanunda, J. G. O. Tepper (SAMA); Tintinarra Lesron, 1993,GLH (SAMA); Toopawarinna Hill, 1994, SANPSDS (SAMA); Venus Bay, 7.5 km NW, 1992, SEG(SAMA); Victor Harbor, 1973, PJMG (ANIC); Victor Harbor, 1990, BBL (SAMA); Victor Harbour, 1969,RHF (SAMA); Vokes Hill Junction, 1994, JAF (SAMA); Waikerie, with Ogyris, 1995, RPF (SAMA);Waikerie, 1995, S. Xuereb, NMVA; Waikerie, with Ogyris, 1992, M. Moore (SAMA); Warburton Ridge, nrKalamurina, 1975, JAF (ANIC); Warooka, with Ogyris, 1995, LHU (SAMA); Weebubbie, 1960, P. Aitken(SAMA); West Avenue Range, 1996, AMA (SAMA); Western Flat, 1994, T. Croft and G. Carpenter(SAMA); William Ck,. Nunna Bore, 1974, JAH (SAMA); Willunga, 1990, RHF (SAMA); Willunga SawPit Rd, 1993, RHF (SAMA); Yalata, 50 km NW, 1984, SANPNS (SAMA); Yalata Roadhouse, 1984,SANPNS (SAMA); Yelpawaralinna, W. H. Warburton Ck., 1993, JAF DH (SAMA); Yerilla Ck nr L.Callabonna, 1953, GFG (SAMA). Tasmania: Cape Barren I., 1949, NBT (SAMA); Cape PortlandSanctuary, 1992, BBL (SAMA); Swansea, 1962, L. Weatherill (ANIC). Victoria: Big Desert, 1973, D. F.Crosby (ANIC); Big Desert, with Ogyris, 1975, AAT (SAMA); Grampians, 10 km S Halls Gap, 1992, AMAand JDE (SAMA); Kaniva, 30 mi S, 1974, BBL (ANIC); Meringur, 23.8 km WSW, 1987, ALY (NMVA);Mildura, 1927, J. C. Myers (SAMA); Mildura, with Ogyris, 1995, RPF (SAMA); Mildura, with Ogyris,1992, AAT (SAMA); Murrayville, 19.8 km SW, 1987, ALY (NMVA); Sea Lake, J. C. Gouldie (SAMA);Wilsons Promontory, 1967, R. D. Howell (ANIC); Wilsons Promontory Cemetry, 1989, S. Morrison(SAMA); Lorne Forest Park, 1995, AMA (SAMA). Western Australia: Banay Ck, with Ogyris, RPF(SAMA); Bannister, 3 km SE, 1992, MAA (SAMA); Beverley, F. H. du Boulay (ANIC); Boonaring Forestnr Williams, 1992, AMA and WMA (SAMA); Bunbury, 1969, BBL (ANIC); Canegrass, 70 km NNWKalgoorlie, 1977, JEF (ANIC); Cape Arid Natl Pk, Seal Creek, 1989, AHB ANA (SAMA); Cape Arid NatlPk, Vokinup Bay, 1989, AHB and ANA (SAMA); Cape LeGrande Cons. Pk, 1994, MAA (SAMA); CapeLeGrande Cons. Pk, Lucky Bay, 1993, AMA and WMA (SAMA); Denmark, 1929, TG (ANIC); Denmark,10 km NE, 1994, MAA (SAMA); Denmark, 20 km NE, 1994, MAA (SAMA); Eneabba, 1987, R. P.McMillam (WAM); Esperance, 1971, BBL (SAMA); Esperance, 1986, AMA (SAMA); Esperance, 11 MileBeach, 1986, AMA (SAMA); Esperance, 30 mi E, 1969, BBL (ANIC); Esperance, 40 km N, 1994, MAA(SAMA); Esperance, 4 mi NE, 1947, TG (ANIC); Esperance, 50 km N, 1994, MAA (SAMA); Esperance, 8mi NW, 1969, RWT (SAMA); Esperance, Lane Rd + Savage Rd, 1986, AMA (SAMA); Esperance, 1977,JEF (ANIC); Esperance Helms Arboretum, 1990, AMA (SAMA); Esperance Lort Ridge, 1990, AMA(SAMA); Esperance Ocean View, 1988, AMA (SAMA); Esperance Wittenoom Hills, 1988, AMA (SAMA);Gold Holes, Stirling Range, 1969, RWT (ANIC); Gora (as Goora) Hill, 1958, TG (ANIC); Grass Patch, 25km W, 1989, AMA (SAMA); Kalgoorlie, 10 km W, 1994, MAA (SAMA); Kalgoorlie, 12 km SW LakeDouglas, with Ogyris, 1991, RPF (SAMA); Karragullen, 1985, P. S. Ward (ANIC); Kau Rock Rd,Quinlivan, 1986, AMA (SAMA); Kings Park, 1993, AMA and WMA (SAMA); Lancelin, 1991, M. S. andB. J. Moulds (AM); Leeman, with Ogyris, 1993, AAT (SAMA); Leeman, RH (SAMA); Mt Arid, CapeLeGrande Natl Pk, with Ogyris, 1991, RPF (SAMA); Mt Ragged, with Ogyris, 1991, RPF (SAMA); MtRagged, 3 mi SbyW, 1969, RWT (ANIC); Mt Arid, 14 mi NWbyW, 1969, RWT (ANIC); Mt Ragged, 1969,BBL (ANIC); Mundaring, JC (ANIC); Mundaring Weir, 1929, TG (ANIC); Norseman, 1974, BBL(SAMA); North Bannister, 10 km NW, 1994, MAA (SAMA); North Bannister, 2 km NW, 1994, MAA(SAMA); North Bannister, 55 km toward Wandering, 1993, AMA and WMA (SAMA); Ongerup, 19 mi E,1947, TG (ANIC); Peak Charles, 1993, AMA (SAMA); Peron Peninsula Nanga Neck, 1972, EGM(SAMA); Perth, JC (ANIC); Perth, Kings Park, 1993, AMA and WMA (SAMA); Perth, Kings Park, 1969,BBL (ANIC); Perth, Pickering Brook, 1993, M. R. Williams (SAMA); Pindar, 1963, C. Mercovich (ANIC);Porongurup, 30 km NE, 1994, MAA (SAMA); Ravensthorpe, 1969, BBL (SAMA); Ravensthorpe, 28 miESE, 1947, TG (ANIC); Ravensthorpe, 53 mi SE, 1969, RWT (SAMA); Salmon Gums, 30 km W, 1993,AMA (SAMA); Salmon Gums + Lake King Rd, 1993, AMA and WMA (SAMA); Serpentine Lakes, 1994,JAF (SAMA); Stirling Range, with Ogyris, 1990, AAT (SAMA); Tammin, TG (ANIC); Thomas Ridge, 3km N mouth, 1969, RWT (ANIC); Thomas Ridge, 1958, TG (ANIC); Toolinna, 1973, M. G. Brooker(ANIC); Walpole, 10 km W, 1994, MAA (SAMA); Williams, 1994, MAA (SAMA); Williams, BoonaringForest, 1992, AMA and WMA (SAMA); Youanmi, on Paynes Find Rd, 10 mi S, 1975, A. M. and M. J.Douglas (ANIC); Watheroo, 1993, RH (SAMA).


Diagnosis

Camponotus terebrans is clearly distinguished from all other ants by the followingcharacters. (1) As a formicine, the acidopore in lateral view is tapering to the orifice, which isfringed with short setae. (2) As a member of the genus Camponotus, the antennae are inserted inthe head capsule distant from the clypeus by a distance greater than the diameter of the antennalfossae. (3) As a member of the C. wiederkehri group, it displays, in lateral and ventral view,J-shaped setae attached to ventral mouthparts usually 4–8 in number and about 0.5 mm long(Fig. 7). It could be confused with Melophorus spp., which possess superficially similarJ-shaped setae, but in Melophorus the antennae are inserted close to the clypeus. (4) The node inlateral view has a summit that is angular or sharp and never smoothly convex. (5) Tibiae arepilose, and the longest setae display an uneven inclination ranging from 30° to 70° and areseparated by a distance that is equal to or greater than their length, with shorter, more adpressedsetae, more closely spaced (Fig. 8). (6) Antennal scapes are pilose, the longest setae are nearerect and sparse along the length of the scape, and the shorter setae are more adpressed anddenser.


Fig. 7. Head of minor worker(ventral view) showing closed mandiblesand closed mouthparts and fourcharacteristic J-shaped setae, attached neararrow. Scale line = 500 µm.

Fig. 8. A section of a mid tibia showing upstanding setae.The flexor or inner side is on the left, where four (of the totalnine or 10) short erect bristles, which are much coarser than thesetae, may be seen. Scale line = 500 µm.

Worker Description

Major worker. Lateral view. Head dark brown to red-brown, sides finely reticulate, glossy.Scape dark brown to black. Funiculus dark brown to yellow-brown. Vertex with 6–10 long andshort setae. Gula with 10–20 setae. Dorsal view. Head sides mostly strongly convex thentapering to the front. Vertex straight, concave in some other views. Setae on scape (near centre)distinct, sparse, sub-erect to erect, pubescence adpressed to decumbent, distinct, spaced less thantheir length apart. Frontal carinae, anterior quarter converging then diverging, next quarterstraight diverging, posterior half parallel then diverging. Frontal area, indistinct, small, elongateddiamond shape. Head width reaches maximum near a line through eye centre. Six teeth on themasticatory border plus a small tooth on the basal border near the angle. Clypeus anterior margin(Fig. 9a), lateral quarters transverse straight, median half strongly projecting to the frontbounded by 90° angles, with crenulations. Integument of clypeus finely reticulate, sparsely andfeebly punctate, glossy. Pubescence on clypeus adpressed, indistinct, spaced much greater thantheir length apart. Carina indistinct. Lateral view. Pronotum brown to yellow, sometimesmottled, from an anterior narrow flat plate follows a gentle concavity then a gentle convexity,with 10–20 setae. Mesonotum brown to yellow, sometimes mottled, flatly convex, with 10–20setae. Metanotum distinct (Fig. 10a), usually straight although sometimes convex, sometimessloping down towards the node, usually glabrous, separated from the propodeum by a grooveusually shorter than a quarter of the length of the metanotum, rarely half as long. Propodeumbrown to yellow, with 10–20 setae, dorsum either straight or feebly convex, angle rounded andoften indistinct, generally about 130° but sometimes a little sharper in largest major workers,upper three-quarters of declivity straight, length of dorsal and declining faces about equal.Integument near propodeal spiracle finely reticulate and usually glabrous, pubescence muchdenser below than above, spiracle elongated and placed on the side well forward of declivity.Node brown to yellow, anterior face convex, summit sharp (Fig. 10a), posterior face mostlystraight, pubescence adpressed, usually sparse. Gaster darker than mesosoma, usually black ordark brown, integument of first gastric tergite microscopically finely striate, glossy. Fore coxaalways lighter than pronotum, yellowish rarely whitish. Fore femur always lighter thanpronotum, yellowish rarely whitish. Fore tibia red-yellow to red-brown. Fore tarsus red-yellowto red-brown. Mid-tibiae with outer setae 0.2 mm long, spaced greater than their length apart,inclined about 60°, pubescence adpressed distinct, 9 or 10 bristles in each of 2 rows on innersurface, often hard to isolate from surrounding setae. Front or rear view. Node summit bidentatein largest major workers, otherwise convex with 6–10 long setae.


Fig. 9. Clypeus (anterior margin) and mandibles: (a) major worker and (b) minor worker. Note that theteeth of the minor worker are very blunt and worn, indicating its old age. Scale line = 500 µm.

(a) (b)

Minor worker. Lateral view. Head black or brown to red-brown with sides finely reticulate,glossy. Scape from dark brown to black. Funiculus from dark brown to yellow-brown. Vertexwith 10–50 long and short setae. Gula with 10–20 setae. Dorsal view. Head sides parallel andfeebly convex, vertex strongly convex. Scape (near centre) with distinct sparse sub-erect to erectsetae, pubescence adpressed to decumbent, spaced less than their length apart, distinct. Frontalcarinae overall nearly parallel, anterior quarter feebly converging, centre half straight, feeblydiverging, posterior quarter feebly converging. Frontal area distinct, clypeal margin convex.Head width reaches maximum near eye centre. Six teeth on the masticatory border plus a smalltooth on basal border near the angle. Clypeus anterior margin (Fig. 9b) lateral sixths straighttransverse, median two-thirds evenly convex, strongly projecting. Clypeus finely reticulate,glossy. Pubescence on clypeus decumbent, coarse, sparse. Carina distinct. Lateral view.Pronotum brown to yellow, from an anterior flat plate with short erect setae there is a short stepto the evenly flatly convex dorsum, with 10–30 setae plus decumbent and adpressed pubescence.Mesonotum brown to yellow, flatly convex, with 10–20 setae. Metanotum obsolete (Fig. 10b).Propodeum brown to yellow, with 10–20 setae, dorsum straight, angle about 160° rounded,declivity upper three-quarters straight, ratio dorsum to declivity exceeds 2 in smallest minors.Integument near propodeal spiracle finely reticulate, adpressed pubescence usually partiallyhiding integument, spiracle elongated and well forward of declivity. Node brown to yellow,anterior face convex, summit angular, posterior face lower three-quarters straight, upper quarterconvex, pubescence adpressed, usually sparse. Gaster darker than mesosoma, usually black ordark brown, integument of first gastric tergite microscopically finely striate, glossy. Fore coxausually lighter than pronotum, yellowish rarely whitish. Fore femur usually lighter thanpronotum, yellowish rarely whitish. Fore tibia red-yellow to red-brown. Fore tarsus red-yellowto red-brown. Mid tibia with setae on outside 0.2 mm long, spaced greater than their lengthapart, inclined about 60°, pubescence outside adpressed, distinct, with 2 rows of 9 or 10 bristleson insides. Front or rear view. Node summit strongly convex with 6–10 long setae.

Medium worker. Characters are intermediate between major and minor workers.

Distribution

The known distribution of C. terebrans is shown in Fig. 11 and is confined to patches ofsandy soil in southern Australia.


Fig. 10. Mesosoma and node in lateral view: (a) major worker and (b) minor worker. Scale line = 500 µm.

(a) (b)

Camponotus darlingtoni Wheeler

Camponotus (Mymophyma) darlingtoni Wheeler, 1934: 160.Camponotus (Mymophyma) rottnesti Donisthorpe, 1941: 239 [unnecessary new name: see Bolton (1995)

for details].

Material Examined

84 workers and 1 queen from Margaret R., 14 workers from Rottnest I., 1 worker from Mullewa, 1worker from Kings Park, Perth (MCZ); 3 workers from Rottnest I. (WAM); 3 workers from Margaret R.(NMVA).

Discussion

Brown (1956) synonymised Wheeler’s C. darlingtoni with Smith’s C. testaceipes (as notedabove, C. testaceipes is preoccupied and its replacement name is C. terebrans). This synonymywas based on Dr E. O. Wilson’s comparison of Wheeler’s types in MCZ with Smith’s type inBMNH, as Dr Brown did not compare the specimens himself. During this study, we havere-examined this material and find that C. darlingtoni is a valid species distinct fromC. terebrans. In particular, C. darlingtoni lacks the cluster of elongate J-shaped setae on themouthparts, a characteristic of the C. wiederkehri species-group, to which C. terebrans belongs.Thus, C. darlingtoni is removed from synonymy with C. terebrans and is here excluded fromthe C. wiederkehri species-group. Unfortunately, the exact placement of this species will requirea broader examination of the rich and diverse Australian Camponotus fauna and is beyond thescope of this paper. To secure the identity of C. darlingtoni, a worker from Margaret River,Western Australia, which is housed in the MCZ collection, is here designated a lectotype.


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Fig. 11. Known distribution of Camponotus terebrans.

Biology

Camponotus terebrans is confined to sandy soils. It can show a very patchy distribution in theGawler Ranges and Sturts Stony Desert, where patches of sand may be only a few hectares. Thesesandy areas are generally dominated by C. terebrans while the surrounding heavier soils arepopulated by a different assemblage of ants. For example, at Scrubby Peak (32°378S, 135°118E)in the Gawler Ranges, South Australia, in a deposit of sand, C. terebrans was collected, whereasin the surrounding heavier soil, Camponotus gouldianus Forel, Camponotus loweryi McArthur,Camponotus wiederkehri Forel and Camponotus consobrinus Erichson were collected.

The species is notable for its ability to quickly colonise disturbed habitats. For example, inHelm’s Arboretum near Esperance, Western Australia (33°348S, 121°538E), C. terebrans wasobserved nesting in the sandy soil that had been raised up by a grader during the construction ofa track across a swamp. At Mount Hope, South Australia (37°298S, 140°108E), C. terebrans wasone of the first ants to colonise a sandy site after it was stabilised in 1948 following 50 years ofdevastation by rabbits, by which time there was no vegetation left and most of the sandy topsoilhad blown away. Within a year or two of stabilisation, grasses and trees returned, as didC. terebrans to become the dominant ant. The first colonies were established beneath stones andstumps, but as Allocasuarina stricta (Macklin) L. Johnson and Melaleuca lanceolata Ottogerminated and increased in size, the ants excavated tunnels close to the trunks to galleriesbelow. Occasionally, trees with a trunk diameter of up to 100 mm could be found with all soilfor about 1 cm surrounding the bark excavated for some depth, causing the tree to be loose in thesoil and sometimes causing its death. Similarly loosening soil around shrubs occurred in agarden at Kandara, near Furner.

The senior author has observed aggressive behaviour by C. terebrans towards two other species.At Mount Hope in 1948, a colony of Iridomyrmex purpureus Sm. occupied the heavier ‘terra rosa’soil on the periphery of the sandy soil, but by 1958 the species had vanished, presumably becauseof competition with C. terebrans. To this day, C. consobrinus still occupies portions of theadjoining ‘terra rosa’ soil surrounding the C. terebrans territory. Fighting between C. terebrans andC. consobrinus has been observed at the boundary in summer early in the morning.

The senior author has also observed C. terebrans in what appears to be a dominant role at asand quarry in Lorne Forest Park, Otway Ranges, Victoria (38°278S, 143°588E), where nests ofants of the Myrmecia pilosula species-group are common. These nests are mounds constructedfrom soil and twigs close to trunks of low bushes. Some were occupied by ants of theM. pilosula species-group while others were occupied by C. terebrans; it appears thatC. terebrans has taken over nests originally constructed by the former.

Camponotus terebrans also appears to be the dominant ant in the bed of Strzelecki Creeknear Yaningurie Waterhole (28°588S, 140°068E), South Australia, where its behaviour isspectacular. Here, the ant was observed to construct long underground tunnels a centimetre ortwo below the surface of the sandy soil. By scraping the soil away from the top of these tunnels,C. terebrans was revealed running to and fro. The tunnels are prominent. They are marked bylong narrow semicircular mounds of soil about 0.5 cm tall on the surface of the creek bed andare often more than 50 m long, connecting colonies near the centre of the creek bed to largeEucalyptus trees on the bank. In July 1989, about 100 of these tunnels were obvious in the creekbed in a space of 5 km downstream from Yaningurie Waterhole, but in July 1994, only threesuch tunnels could be found.

An extensive collection of foraging C. terebrans workers was undertaken at Mount Hope,South Australia. This collection demonstrates that only the smallest workers venture far from thenest. Foragers were collected at night by beating a Banksia marginata Cav. shrub growing about20 m from a nest. A random sample of workers was then collected from this nest. The headwidths and lengths for these ants are plotted in Fig. 12. Only the smallest workers were foundforaging on the Banksia shrub while both small and large workers were found in the nest. Thissuggests that major and medium workers of C. terebrans do not travel far from the nest whileminor workers can forage much more broadly. Where C. terebrans is densely populated, such as


at Mount Hope and Stockyard Plain, diurnal foragers have been observed, although nocturnalforaging is more general. Nests of Camponotus aurocinctus (Smith) have been observed within afew metres of nests of C. terebrans in sandy soil at Stockyard Plain and at DanggaliConservation Park.

Associations with Other Invertebrates

Coleoptera : Tenebrionidae. There is a relationship between C. terebrans and thetenebrionid beetle Camponotiphilus fimbricollis Lea, involving co-habitation (Mathews 1993).C. terebrans and Camponotiphilus fimbricollis have been taken from the same nest at Beverley,Western Australia (32°078S, 116°568E), by F. H. duBoulay (SAMA) and at Hattah Lakes,Victoria (34°458S, 142°208E) (NMVA).

Lepidoptera. Camponotus terebrans is believed to have a mutualistic relationship with abutterfly (Ogyris spp.). Localities at which the larvae of Ogyris spp. and C. terebrans have beencollected from the same nest are recorded in the ‘Other Material Examined’ section as ‘withOgyris’. In these cases, the entrance to the nest was generally close to a food plant of Ogyris,such as Choretrum glomeratum R. Br.

Conclusion

Prior to this study, what is here considered to be C. terebrans was in a state of someuncertainty, with the existence of several valid species plus a subspecies. With the establishmentof the new synonyms C. latrunculus victoriensis and C. myoporus, the taxonomy of C. terebransis now underpinned by an unambiguous diagnosis, despite the considerable variation displayedin both morphological and allozyme characters. This study represents useful baseline


Fig. 12. Head width and head length of Camponotus terebrans workers taken at Mount Hope. Individualswere taken when foraging on a shrub, Banksia marginata (n = 65), and dug from a nearby nest (n = 285).There is some overlap in the figure for small head widths.

information that will facilitate the future documentation of morphological, ecological andbehavioural differences that may exist within this variable species. In time, this could help fieldbiologists to further document the natural history of C. terebrans.

Acknowledgments

This work has been made possible by grants from ABRS. In addition, we are extremelygrateful for the use of facilities and support provided by the South Australian Museum and forthe help given by Mrs M. Anthony, Dr C. H. S. Watts, Mrs J. Forrest OAM, Dr E. G. Mathews,Miss J. Thurmer, Mrs C. Horne, Mr M. Krieg, Dr P. Kolesek, Dr P. J. M. Greenslade and Dr R.W. Taylor (ANIC); Miss Catriona McPhee, Mr Frank Walker and Dr A. L. Yen (NVMA); Dr T.Houston (WAM); Mr W. M. McArthur, Rev. B. B. Lowery SJ (deceased), Mr R. H. Mew andMr Barry Bolton (BMNH); Mr S. P. Cover (MCZ); and two anonymous reviewers.

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Manuscript received 28 April 1997; revised and accepted 17 November 1997


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