Zootaxa 3955 (2): 283–290 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3955.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:97FCBEF2-E95E-47A8-A959-408241A2574D A review of the Myrmecorhynchus (: Formicidae)

S.O. SHATTUCK ARC Centre of Excellence in Vision Science, Research School of Biology, The Australian National University, Building 46, Biology Place, Canberra, Australian Capital Territory 2601, and Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA

Abstract

The Australian endemic ant genus Myrmecorhynchus is reviewed. The genus is known from three species (M. carteri Clark, M. emeryi André and M. nitidus Clark) which are restricted to eastern and southern Australia. Myrmecorhynchus musgravei Clark and M. rufithorax Clark are newly synonymised with M. emeryi André. All species are found in forested areas where they nest arboreally or, less commonly, in soil. Foraging occurs primarily on vegetation and tree trunks.

Key words: Myrmecorhynchus, Australia, , Formicidae

Introduction

Myrmecorhynchus is an endemic Australian genus, known from three species. They occur in forested areas ranging from mallee through rainforest across eastern and southern Australia. All three species are sympatric in Victoria and , with M. emeryi extending westward to south-western Western Australia and northward to central , and with M. carteri occurring in Tasmania (Fig. 1). They are small and inconspicuous and are most often encountered while foraging on vegetation or tree trunks (Fig. 2). Nests are in branches, twigs and vines on shrubs or trees, or in soil. Although they can be locally common they are often overlooked because of their slow movements and arboreal habits. Distribution patterns of Myrmecorhynchus species were examined using the DIVA-GIS software package. Predicted distributions were generated using Domain modelling while species richness was based on the individual species models and calculated using sum set to present/absent where present > 90. The modelling suggests that one of the species, M. emeryi, is likely to be more widely dispersed than currently available material suggests, while the remaining two species appear to be known from throughout their potential range. This is as might be expected for a relatively little-collected group, in this case likely because of their arboreal habits, and additional targeted collecting will likely fill the apparent gaps in our understanding of the distribution of these ants.

Myrmecorhynchus André

Myrmecorhynchus André, 1896: 253. Type-species: Myrmecorhynchus emeryi, by monotypy.

Diagnosis. Mandibles usually with 10 to 13 teeth, although some large individuals with as few as 6. Frontal carinae distinctly arched. Upper surface of mesosoma between metanotum and propodeum always low and flat or concave and never expanded upwards. Worker caste variable in size (strongly polymorphic) with distinct major and minor workers. Smaller workers of Myrmecorhynchus are identifiable by the large number of mandibular teeth. However larger workers have a reduced number of teeth (down to 6 in some individuals) and are similar to some species of Notoncus. These individuals can be identified by the configuration of the frontal carinae (curved in Myrmecorhynchus, straight in Notoncus) and the polymorphic worker caste with distinct majors and minors

Accepted by J. Longino: 13 Apr. 2015; published: 4 May 2015 283 (Notoncus is weakly polymorphic and without distinct majors and minors). Additionally, some species of Notoncus have an upwards projection on the upper surface of the mesosoma at the metanotal groove. This region of the mesosoma is always flat or concave in Myrmecorhynchus.

FIGURE 1. Predicted distribution and species richness of the genus Myrmecorhynchus based on range models of individual species. Species are absent from white areas, a single species is predicted to occur in yellow areas, two species are predicted from orange areas, and three species from red areas.

List of species carteri Clark emeryi André musgravei Clark n. syn. rufithorax Clark n. syn. nitidus Clark

Key to Myrmecorhynchus species based on workers

1. Mesosomal dorsum and legs with abundant erect hairs ...... emeryi -. Mesosomal dorsum with at most a few scattered erect hairs, legs with appressed pubescence but lacking erect hairs...... 2 2. Extensive yellow on clypeus covering at least 1/3 of surface; mandibles completely yellow (except dark teeth); propodeum rel- atively low and flat, metapleural groove very shallow (Fig. 6) ...... nitidus -. At most a narrow band of yellow on anterolateral clypeal margin; mandibles bicoloured, yellow basally and dark apically; propodeum higher and more rounded, metapleural groove more strongly angled (Fig. 3) ...... carteri

284 · Zootaxa 3955 (2) © 2015 Magnolia Press SHATTUCK FIGURE 2. Myrmecorhynchus emeryi worker, 9.5km S Bungendore, New South Wales, S.O. Shattuck (ANIC32-042179).

Myrmecorhynchus carteri Clark (Fig. 3)

Myrmecorhynchus carteri Clark, 1934: 43.

Types. Syntype workers from Barrington Tops, New South Wales (H. J. Carter) [Museum Victoria, Melbourne, examined] and Kinglake, Victoria (T. Greaves) [location unknown]. Diagnosis. Mesosomal dorsum with at most a few scattered erect hairs, legs with appressed pubescence but lacking erect hairs; at most a narrow band of yellow on anterolateral clypeal margin; mandibles bicoloured, yellow basally and dark apically; propodeum higher and more rounded and metapleural groove more strongly angled when compared to M. nitidus. Comments. Myrmecorhynchus carteri occurs from north-coastal New South Wales south to Tasmania (it is the only species known from Tasmania). It is found in mallee, thick scrub, dry to wet sclerophyll (including Eucalyptus pauciflora, E. perriniana and E. stellulata forests), coastal scrub (wet scrub; hillside, black soil; at water’s edge) and rainforest. Nests are in dead branches or twigs above the ground (including Acacia sophorae, Eucalyptus perriniana and E. stellulata) or occasionally on the ground, or less commonly in soil under rocks. One nest was found in a dead, rotten branch with a colony of M. emeryi; however the relationship between these ants, if any, is unknown. Foragers are commonly found on tree trunks and are known to visit Eucryphia lucida flowers and wander into malaise traps. The larvae were described by Wheeler & Wheeler (1970). Morphologically this species is similar to M. nitidus, differing in colour pattern and the shape of the metanotal groove and propodeum. While colour pattern shows considerable variation in M. emeryi, this does not seem to be the case here. All known specimens show either the M. carteri or M. nitidus patterns with no intermediate individuals being found. Thus colour suggests that these two taxa are distinct. The shape of the metanotal groove and propodeum also support this conclusion. All known specimens of M. carteri have the propodeum higher and more rounded dorsally compared to M. nitidus workers, resulting in the metanotal groove being more distinct and angular. As with colour, all specimens show one or the other pattern with intermediate forms unknown. Combined, these characters suggest that two separate taxa are involved rather than a single variable species.

REVIEW OF MYRMECORHYNCHUS Zootaxa 3955 (2) © 2015 Magnolia Press · 285 FIGURE 3. Myrmecorhynchus carteri, minor worker, Moppy Lookout, Barrington Tops State Forest, New South Wales, T.A. Weir (ANIC32-010761).

Material examined. Australia: Australian Capital Territory: Woods Reserve, nr. Gibraltar Falls (Naumann,I.D.). New South Wales: Belmore Falls (Top) (Liepa,Z.); Blue Knob, Nightcap Range (Lowery,B.B.); Dainers Gap (Morrow,P.); Dilgry River Barrington Tops S.F. (Weir,T.A.); East Boyd State Forest, Goanna Rd., 56km SE Bombala (Lambkin,C. & Starick,N.); Kosciuszko National Park, 1.7km ENE of Thredbo (Lambkin,C., Starwick,N. & Recsei,J.); Moppy Lookout Barrington Tops S.F. (Weir,T.A.); Mt. Wilson, Blue Mountains (Horning,D.S., Jr.); New England Nat. Park. via Ebor (Monteith,G.). Tasmania: 4km N Montagu (Lowery,B.B.); 5km E Low Head (Lowery,B.B.); 5km N Emita (Lowery,B.B.); Blythe Sdg. (Greaves,T.); Darling Range, Flinders Island (Lowery,B.B.); Derwent Bridge, Bronte (collector unknown); Devonport (Lowery,B.B.); Friendly Beaches, Freycinet Natl. Pk. (Lowery,B.B.); Greens Beach, Asbestos Range Natl. Pk. (Lowery,B.B.); Headwaters, Olga R., E of Mt. Eleanor (Howard,C., et al.); Leith (Lowery,B.B.); Lower Gordon River, E of Goulds Landing (collector unknown); near Lake Chisholm (Ettershank,G. & Ettershank,J.A.); Olga R. Top Plain, Transects 12 & 13 (Howard,C. & Johnson,C.); Sisters Beach (Lowery,B.B.); Thule Rd., Flinders Island (Lowery,B.B.); Upr. Gordon River, N and W of Denison Camp (Howard,C., et al.); Upr. Gordon River, nr. Angel Cliffs (Howard,C., et al.). Victoria: Coopracambra NP, 27km NNE Cann River, Beehive Creek (Lambkin,C. & Starick,N.); Ferntree Gully (Lowery,B.B.); Flinders (Lowery,B.B.); Harrietville (Lowery,B.B.); Kinglake (Greaves,T.); Leongatha (Lowery,B.B.); Mt. Dandenong (Schwarz,M.); Woori Yallock (Thorn,L.B.).

286 · Zootaxa 3955 (2) © 2015 Magnolia Press SHATTUCK Myrmecorhynchus emeryi André (Figs 2, 4, 5)

Myrmecorhynchus emeryi André, 1896: 254. Myrmecorhynchus musgravei Clark, 1934: 43. NEW SYNONYM Myrmecorhynchus rufithorax Clark, 1934: 46. NEW SYNONYM

Types. M. emeryi: Holotype worker from the Victorian Alps, Victoria (Musee National d'Histoire Naturelle, Paris, examined). M. musgravei: Holotype male from Lamington Nat. Park (as National Park), Queensland (A. Musgrave, 19 December 1926) (Australian Museum, Sydney). M. rufithorax: Syntype workers from Warburton, Victoria (L. B. Thorn, in trees) (Museum Victoria, Melbourne). Diagnosis. Mesosomal dorsum and legs with abundant erect hairs. Body colour variable, ranging from reddish to reddish-brown.

FIGURE 4. Myrmecorhynchus emeryi, major worker, Gerroa, New South Wales, B.B. Lowery (ANIC 32-010956).

Comments. Myrmecorhynchus emeryi is the most widely distributed species in the genus, occurring from south-eastern Queensland (with a single record further north) south through New South Wales and Victoria to south-western Western Australia. It is found in a range of habitats including mallee, sandstone scrub and moist

REVIEW OF MYRMECORHYNCHUS Zootaxa 3955 (2) © 2015 Magnolia Press · 287 gully woodlands, dry to wet sclerophyll and rainforest. Nests are found in dead branches, twigs and vines, in soil most commonly at the base of tree but also under rocks or occasionally in moss. Nests have also been found in dead stumps (including a Xanthorrhoea stump). Three females were found in a cavity in a small dead twig 2.5m (8 ft) above the ground. This would suggest that this species is pleometrotic, however additional observations are needed to confirm this. Workers are most commonly seen foraging on trees and low vegetation and have been found feeding at extra-floral nectaries of Acacia pycnantha.

FIGURE 5. Myrmecorhynchus emeryi, minor worker, Mt. Remarkable, Flinders Ranges, South Australia, P.J.M. Greenslade (ANIC32-010964).

When describing the taxon M. rufithorax, Clark (1934) commented that this species was “near M. emeryi Andre”, but gave no specific characters to separate these two taxa. With the presently available material, there seems little doubt that M. rufithorax represents the pale form of a single variable species, with M. emeryi representing the darker form. Intermediately coloured individuals are abundant and no additional characters could be found to suggest that more than a single species is involved. Because of this these two taxa are here considered synonyms. Myrmecorhynchus musgravei, also described by Clark (1934), was based on an unassociated male. Clark mentions several characters which separate M. musgravei from M. emeryi, including differences in colour (testaceous in M. musgravei, reddish yellow in M. emeryi) and wing morphology (discoidal cell large in M. musgravei, small in M. emeryi). However the limited available male material makes it difficult to assess the taxonomic significance of these and other characters used by Clark (1934). Numerous worker collections from the region of the type locality of M. musgravei in south-eastern Queensland suggest that a single species occurs there. There is no evidence for two distinct species in this region. Combined, the presently available information suggests that M. musgravei is the male of M. emeryi and not a separate species. The major worker of this species was described by Emery (1898), the queen and male by Clark (1934) and the larva by Wheeler (1917) and Wheeler & Wheeler (1968).

288 · Zootaxa 3955 (2) © 2015 Magnolia Press SHATTUCK Material examined. Australia: Australian Capital Territory: Black Mountain (Greaves,T.); Black Mountain Reserve (Liepa,Z.); Black Mt. (Greaves,T.; Lowery,B.B.); Black Mt., Canberra (Lowery,B.B.); Mount Ainslie (Lowery,B.B.). New South Wales: 10mi. S Kiama (Lowery,B.B.); 20km E Legume (Greenslade,P.J.M.); 4 mi. E of Berry (Lowery,B.B.); 9.5km S Bungendore (Shattuck,S.O.); Berry Mountain, Berry (Lowery,B.B.); Bilambil (Lowery,B.B.); Blue Knob Mt., Nightcap Range (Lowery,B.B.); Bogan River (Armstrong,J.); Bola Creek NP, Sydney (Lowery,B.B.); Cambewarra Mountain (Lowery,B.B.); Cool and Candle Crk., Kuring-gai Chase (Lowery,B.B.); Cowan, Sydney (Lowery,B.B.); Dalrymple Forest, Pymble, Sydney (Lowery,B.B.); Foxground (Lowery,B.B.); Frenchs Forest, Sydney (Lowery,B.B.); Gerroa (Lowery,B.B.); Gerroa, 10mi. S Kiama (Lowery,B.B.); Gibraltar Ra. Nat. Pk. via Glen Innes (Monteith,G.); Hills to W of Mudgee (Lowery,B.B.); Jerusalem Bay, Cowan (Lowery,B.B.); Kyogle (Froggatt,W.W.); Mt. Saddleback, Kiama (Lowery,B.B.); Mt. Warning (Lowery,B.B.); Riverview College, Lane Cove, Sydney (Lowery,B.B.); Riverview College, Sydney (Lowery,B.B.); Runnyford Ck., Nelligen (Liepa,Z.); St. Ives, Frenchs Forest, Sydney (Lowery,B.B.); Tooloom Plateau, 14km W Urbenville (Naumann,I.D.); W of Mt. Warning (Lowery,B.B.); Wallerawang (Lowery,B.B.); Wentworth Falls (Wheeler,W.M.); Wentworth Falls, Blue Mts. (Lowery,B.B.). Queensland: 13km NE Eungella (Ward,P.S.); 1km SSE Eungella (Ward,P.S.); 50mi. W Mackay, Eungella NP (Lowery,B.B.); 7km NNW North Tamborine (Ward,P.S.); Diggings Road, Eungella (Monteith,G.B. & Thompson,G.); Lamington N.P. (Andersen,A.N.); Mt. Tamborine (Greaves,T.); Palm Grove N.P., 8km ESE North Tamborine (Shattuck,S.O.); Tomewin (Lowery,B.B.); Tomewin, (Lowery,B.B.). South Australia: Belair Rec. Park, Adelaide (O'Dowd,D.J.); Bridgewater (Greenslade,P.J.M. & Hutson); Kangaroo Is., Breakneck R. (Greenslade,P.J.M.); Mt. Bold Reservoir, SE of Adelaide (Ward,P.S.); Mt. Lofty (Lowery,B.B.); Mt. Lofty summit (Gross,G.F.); Mt. Remarkable, Flinders Ranges (Greenslade,P.J.M.); Sevenhill (Lowery,B.B.); Spring Gully (Lowery,B.B.); Streaky Bay (Lowery,B.B.); Upper Sturt (Herridge,J.A.); Worlds End, 23km S Burra (Lowery,B.B.). Victoria: 15km WNW Yaapeet, Wyperfeld Natl. Pk (Andersen,A.N.); 20km S Ouyen (Shattuck,S.O.); Harrietville (Lowery,B.B.); Heathcote (Lowery,B.B.); Hurstbridge (Lowery,B.B.); Leongatha (Lowery,B.B.); Warrandyte (Greaves,T.); Woori Yallock (as Wooriyalock) (Thorn,L.B.). Western Australia: 11 mi. E Pingrup (Greaves,T.); 15km N of Norseman (Lowery,B.B.); Corrigin (Lowery,B.B.); Grass Patch (Lowery,B.B.); Junana Rock, 9km NW Mt. Ragged (Taylor,R.W.); Pingrup (Greaves,T.); Tingledale (Taylor,R.W.).

Myrmecorhynchus nitidus Clark (Fig. 6)

Myrmecorhynchus nitidus Clark, 1934: 44.

Types. Syntype workers, queens and males from Cheltenham, Victoria (L. B. Thorn, in dead branch of tree) [4 workers in Australian National Collection, Canberra, examined; 1 worker in Museum Victoria, Melbourne, examined] and workers and males from Canberra, ACT (T. Greaves, in limb of tree, 20m (70 ft) above ground) [2 workers in Australian National Insect Collection, Canberra, examined; 1 worker in The Natural History Museum, London, examined]. Diagnosis. Mesosomal dorsum with at most a few scattered erect hairs, legs with appressed pubescence but lacking erect hairs; extensive yellow on clypeus covering at least 1/3 of surface; mandibles completely yellow (except dark teeth); propodeum relatively low and flat and metapleural groove very shallow when compared to M. carteri. Comments. This is the least common species of Myrmecorhyncus, being known from only a handful of specimens from the vicinities of Melbourne and Canberra. Essentially nothing is known concerning the biology of this species. For notes on the taxonomic status of this species as distinct from the closely related M. carteri, see Comments under that species above. Material examined. Australia: Australian Capital Territory: Canberra (Greaves,T.); Lees Creek (as Lees Springs) (collector unknown). Victoria: Cheltenham (Thorn,L.B.); Xavier College, Kew (Lowery,B.B.).

REVIEW OF MYRMECORHYNCHUS Zootaxa 3955 (2) © 2015 Magnolia Press · 289 FIGURE 6. Myrmecorhynchus nitidus, minor worker, Xavier College, Kew, Victoria (ANIC 32-066442).

Acknowledgements

This project was initiated with support from CSIRO and completed with funds provided by the Australian National University and Harvard University (through E.O. Wilson and the Arthur L. Green Fund).

References

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