Integrative taxonomy reveals cryptic diversity in North American Lasius ants, and an overlooked introduced species

Biological invasions are a grave threat to ecosystems. The black garden ant (Lasius niger) is a pest species in Europe. Current literature states that L. niger occupies a disjunct native distribution in the Holarctic, however, based on recent work, we re-evaluate this distribution. The native range of L. niger is reconsidered based on phylogenetic relationships (nine mitochondrial and nuclear markers, 5670 bp), DNA-barcoding (98 Holarctic specimens), morphometry (88 Holarctic specimens, 19 different measurements) and subjective assessment of phenotype. The potential spread of this species is estimated using ecological niche modeling. Lasius niger is more closely related to other Palearctic species than to the Nearctic ants known under this name. The latter are described as a distinct species, L. ponderosae sp. nov. However, DNA-barcoding discovered established populations of L. niger in metropolitan areas in Canada (Vancouver and Halifax). We describe a morphometrical method to delineate L. ponderosae sp. nov. and L. niger. MtDNA diversity and divergence is high within L. ponderosae sp. nov., but low within L. niger. More than 1,000,000 km 2 are suitable as a habitat for L. niger in North America. This case emphasizes the critical role of integrative taxonomy to detect cryptic species and identify potential biological invasions in their nascent stages

Desert springs: deep phylogeographic structure in an ancient endemic crustacean (Phreatomerus latipes)

Extent: 13p.Desert mound springs of the Great Artesian Basin in central Australia maintain an endemic fauna that have historically been considered ubiquitous throughout all of the springs. Recent studies, however, have shown that several endemic invertebrate species are genetically highly structured and contain previously unrecognised species, suggesting that individuals may be geographically ‘stranded in desert islands’. Here we further tested the generality of this hypothesis by conducting genetic analyses of the obligate aquatic phreatoicid isopod Phreatomerus latipes. Phylogenetic and phylogeographic relationships amongst P. latipes individuals were examined using a multilocus approach comprising allozymes and mtDNA sequence data. From the Lake Eyre region in South Australia we collected data for 476 individuals from 69 springs for the mtDNA gene COI; in addition, allozyme electrophoresis was conducted on 331 individuals from 19 sites for 25 putative loci. Phylogenetic and population genetic analyses showed three major clades in both allozyme and mtDNA data, with a further nine mtDNA sub-clades, largely supported by the allozymes. Generally, each of these sub-clades was concordant with a traditional geographic grouping known as spring complexes. We observed a coalescent time between ~ 2–15 million years ago for haplotypes within each of the nine mtDNA sub-clades, whilst an older total time to coalescence (>15 mya) was observed for the three major clades. Overall we observed that multiple layers of phylogeographic history are exemplified by Phreatomerus, suggesting that major climate events and their impact on the landscape have shaped the observed high levels of diversity and endemism. Our results show that this genus reflects a diverse fauna that existed during the early Miocene and appears to have been regionally restricted. Subsequent aridification events have led to substantial contraction of the original habitat, possibly over repeated Pleistocene ice age cycles, with P. latipes populations becoming restricted in the distribution to desert springs.Michelle T. Guzik, Mark A. Adams, Nicholas P. Murphy, Steven J.B. Cooper and Andrew D. Austi

Aphaenogaster poultoni Crawley

Aphaenogaster poultoni Crawley (Figs 6, 13, 14, 19, 20, 29) Aphaenogaster poultoni Crawley, 1922 b: 17. Types. Worker syntypes (10 in WAMP, examined) from Beenup [?=Beenyup Brook or Beenyup Swamp], Western Australia; 1 worker syntype without locality data in MVMA. Diagnosis. Majority of hairs on venter of head located laterally and forming a distinct psammophore (Fig. 14); eye relatively small (EI less than 21, Fig. 19); scape relatively short (SI less than 106, Fig. 20); mandibular sculpture composed of irregularly sized striations (Fig. 6); petiolar node (in dorsal view) wider than long. This species is most similar to A. barbigula. These two species can be separated by the pattern of sculpturing on the mandibles and the shape of the petiolar node. There is a tendency for the head of A. poultoni to be more square (when viewed in full face view, the lateral and posterior margins are separated by an angle) compared to both A. barbigula and A. mediterrae (in which the head is essentially uniformly arched behind the eyes in full face view). However, all species show moderate variation in the shape of the head with essentially identical morphologies being found in some individuals of all three species. Thus while this character is indicative of A. poultoni it is not diagnostic. Description. Posterior margin of head nearly flat in full face view, extending laterally of the occipital collar before passing through a distinct posterolateral corner into the lateral margin of the head. Majority of hairs on venter of head located laterally and forming a distinct psammophore, only scattered hairs on central portion. Mandibular sculpture composed of irregularly sized striations. Erect hairs on mesosomal dorsum tapering to sharp points. Propodeal spines reduced to small denticles or sharp angles. Petiolar node (in dorsal view) wider than long; Measurements. Worker (n = 11). CI 86–98; EI 16–21; EL 0.16–0.22; HL 0.93–1.28; HW 0.85–1.18; ML 1.30–1.86; MTL 0.68 –1.00; SI 90–105; SL 0.85–1.16. Material examined (in ANIC unless otherwise noted). Western Australia: 100km E Norseman (Lowery,B.B.); 10km S Mullewa (Lowery,B.B.); 10mi. W Mullewa (Riek,E.F.); 160km ENE Esperance (Ward,P.S.) (ANIC, PSWC); 180km N Geraldton, nr. Billabong Roadhouse (Lowery,B.B.); 20km S Norseman (Lowery,B.B.); 20km W York (Lowery,B.B.); 26mi. NWbyW Norseman (Taylor,R.W.); 3.2km SSW Dongara (Feehan,J.E.); Beenup [Beenyup] (Clark,J.); Beyerley (du Boulay,F.H., MCZC); Brookton Hwy, 20km E Boulders (Lowery,B.B.); Bungulla (Greaves,T.); Caron (Darlington,P.J., MCZC); Darlington (Glauert,G., MCZC); Geraldton (Lowery,B.B.; Mercovich,C.T.; Weatherill; Wheeler,W.M. (ANIC, MCZC); Gayamin Pool, Lower Chittering (Wilson,E.O. and Douglas,A., MCZC); Israelite Bay (T.G.); Kalbarri Nat. Park (Lowery,B.B.); Kings Park, Perth (Clark,J., Lowery,B.B.) (ANIC, MCZC); Moora (Lowery,B.B.); Mullewa (Wheeler,W.M., MCZC); National Park, Darling Range (Wheeler,W.M., MCZC); Perth (Clark,J.; Greaves,T.; Keirath,A.R.) (ANIC, MCZC); Perth, Kings Park (Lowery,B.B.); Rockingham (Glauert,L., MCZC); Toodyay (Lowery,B.B.); Toolinna (Brooker,M.G.); Woongondy, 300mi. N Perth (Mercovich,C.T.). Comments. This species is restricted to a narrow semi-arid band across south-western Western Australia (Fig. 29) and is sympatric with A. mediterrae in south-central Western Australia (see Fig. 28). It is found in coastal scrub, Jarrah forests, dry sandy sclerophyll, mulga woodlands and mallee. Nests are in soil generally with a large crater at the entrance. An exceptionally large mating swarm was noted by McMillan (1977) (as A. barbigula).Published as part of Shattuck, Steven O, 2008, Australian ants of the genus Aphaenogaster (Hymenoptera: Formicidae), pp. 25-45 in Zootaxa 1677 on pages 40-41, DOI: 10.5281/zenodo.18032

Mesostruma spinosa Shattuck, 2007, new species

Mesostruma spinosa new species (Figs 9–12) Description: Holotype worker. TL 2.9, HL 0.69, HW 0.56, CI 81, MandL 0.21, MandI 30, SL 0.41, SI 73, PW 0.40, ML 0.77. In dorsal view the anterolateral corners of pronotum rounded. Dorsum of mesosoma with scattered, shallow foveolate punctures spaced more than their width apart, the area between punctures smooth and lacking sculpturing. Propodeum armed with elongate spines, propodeal lamellae reduced to thin bands and essentially absent. Sculpturing on metapleural gland bulb consisting of indistinct rugae and punctures. Lateral surfaces of postpetiole rounded. First gastral segment immediately posterior of postpetiole smooth. Body colour yellow-red with light infuscation on dorsum of head above eyes, clypeus and first gastral tergite. Material examined: Holotype worker from Western Australia, Manjimup (34 ° 15 'S 116 °09'E), July, 1977 (J. D. Majer) (ANIC, No. 32 -007281). Comments: Mesostruma spinosa can be separated from all other known species in this genus by the presence of distinct, elongate propodeal spines, the lack of broad propodeal lamellae and the narrower head (CI = 81, CI is greater than 88 in the other known species). There is little chance it will be confused with any other species. This species, together with M. inornata and M. loweryi, have only been collected once. Only one species, M. eccentrica, is at all common in Western Australia, having been collected over 10 times. It is likely that additional species remain to be discovered in this part of Australia. The single known specimen of M. spinosa was collected from a pitfall trap. The following modifications to the key provided by Shattuck (2000) will allow the identification of this species. 3. Dorsum of alitrunk with dense, shallow foveolate punctures which are generally spaced less than their width apart, and with the area between the punctures with weak but distinct sculpturing...................... 4 Dorsum of alitrunk with scattered, shallow foveolate punctures which are spaced more than their width apart, the area between the punctures smooth and lacking sculpturing................................................ 4 A 4 A. Propodeum armed with elongate spines, the lamellae reduced to thin bands which are only slightly raised above the underlying propodeal surface ............................................................................. spinosa Propodeum lacking spines and with thick lamellae................................................................................ 5Published as part of Shattuck, Steven O., 2007, New species of Myrmicine ants from Western Australia (Hymenoptera: Formicidae), pp. 47-53 in Zootaxa 1661 on pages 51-52, DOI: 10.5281/zenodo.16319

Aenictus Shuckard 1840

Aenictus Shuckard, 1840 Aenictus Shuckard, 1840: 266. Typhlatta Smith, 1857: 79 (synonym of Aenictus by Forel, 1890: ciii; removed from synonymy as subgenus of Aenictus by Wheeler, 1930: 198; synonym of Aenictus by Wilson, 1964: 444). Type species. Aenictus: Aenictus ambiguus Shuckard, 1860, by original designation. Typhlatta: Typhlatta laeviceps Smith, by monotypy. Diagnosis. Workers of Aenictus may be separated from other Australian ants by their moderately small size (less than about 4 mm), lack of eyes, long slender bodies and long legs. They are superficially similar to some myrmicines but differ in lacking the frontal lobes and in having the antennal sockets completely visible when viewed from the front (myrmicines have frontal lobes that are expanded towards the sides of the head and partly cover the antennal sockets). Some of the smaller, paler species are also similar to Leptanilla workers, but differ in being larger and only ten segments in the antennae rather than 12, and lacking a flexible promesonotal suture. Males of Aenictus can be separated from those of other Australian ants by the exposed antennal sockets and lack of a postpetiole (the gaster is smooth and lacks a constriction between the first and second segments).Published as part of Shattuck, Steven O., 2008, Review of the ant genus Aenictus (Hymenoptera: Formicidae) in Australia with notes on A. ceylonicus (Mayr), pp. 1-19 in Zootaxa 1926 on page 3, DOI: 10.5281/zenodo.18481

Aphaenogaster barbigula Wheeler

Aphaenogaster barbigula Wheeler (Figs 3 –5, 19, 20, 25) Aphaenogaster (Nystalomyrma) barbigula Wheeler, 1916: 221. Types. Lectotype worker (here designated) from Sea Lake, Victoria, G. C. Goudie (MCZC); paralectotypes as follows: Yanco (6 workers, MCZC, examined), New South Wales; Adelaide (3 workers, MCZC, examined), Meningie (5 workers, MCZC, examined); Gawler and Karoonda to Peebinga, South Australia; Dongara (as Dongarra), Gooseberry Hill, Wallaby Island and Beverley, Western Australia; Sea Lake (16 workers, MCZC, examined; 1 worker, MVMA, examined), Victoria. Diagnosis. Majority of hairs on venter of head located laterally and forming a distinct psammophore (Fig. 4); eye relatively small (EI less than 21, Fig. 19); scape relatively short (SI less than 106, Fig. 20); mandibular sculpture composed of regularly sized striations (Fig. 5); petiolar node (in dorsal view) approximately square. The presence of a psammophore will separate this species from all others except A. mediterrae and A. poultoni. It can be separated from these species as outlined in the key above. Description. Posterior margin of head nearly flat in full face view, extending laterally of the occipital collar before passing through a distinct posterolateral corner into the lateral margin of the head. Majority of hairs on venter of head located laterally and forming a distinct psammophore, only scattered hairs on central portion. Mandibular sculpture composed of regularly sized striations. Erect hairs on mesosomal dorsum tapering to sharp points. Propodeal spines reduced to small denticles, or sometimes essentially absent. Petiolar node (in dorsal view) approximately square. Measurements. Worker (n = 8). CI 84–92; EI 22–27; EL 0.23–0.27; HL 1.12–1.26; HW 0.97–1.16; ML 1.58–1.84; MTL 0.96–1.08; SI 104–115; SL 1.11–1.25. Material examined (in ANIC unless otherwise noted). New South Wales: 12km S Coombah (Ward,P.S.) (ANIC, PSWC); 4mi. N Condobolin (Lowery,B.B.); 60mi. S Broken Hill (Lowery,B.B.); 7mi. S Hillston (Lowery,B.B.); Berrigan SF (Lowery,B.B.); Cocoparra NP, nr Griffith (Lowery,B.B.); Emu Vale, nr. Deniliquin (Greenslade,P.J.M. & Valentine,I.); Griffith, bank of Lake Wyangan (Lowery,B.B.); nr. Darling R., Wentworth (Lowery,B.B.); Lake Popilta [Popiltah Lake] (Lowery,B.B.). Queensland: 15 km S by W of Charleville (Upton,M.S.); 30km W St. George (Lowery,B.B.); Beralga c. 50mi. NW St. George (Lowery,B.B.); St. George (Lowery,B.B.); St. George nr. Balonne R. (Lowery,B.B.). South Australia: 15km NW Renmark (Shattuck,S.O.); 18km NNE Meninjie (Greenslade,P.J.M.); 30mi. S Whyalla, Moonabie Ra. (Haycraft,J., MCZC); 32km N Renmark (Shattuck,S.O.); 40km S Oodnadatta (Lowery,B.B.); 5mi. W Tintinara (Greaves,T.); Blythe, 80mi. N Adelaide (Lowery,B.B.); Ferries McDonald Natl. Pk. (Shattuck,S.O.); Gawler Ra., Saline to SE (Greenslade,P.J.M.); Hambridge NP [Hambidge NP] (Britton,E.B.); Kokatha, Gawler Ranges (Greenslade,P.J.M.); Lochiel (Lowery,B.B.); Perponda (Greenslade,P.J.M.); Tailem Bend (Lowery,B.B.); Two Wells (Lowery,B.B.); Victoria Desert, 53km E Vokes Hill (Greenslade,P.J.M.); Warooka, Yorke Peninsula (Lowery,B.B.); Yorke Pen., Hardwicke Bay (Greenslade,P.J.M.). Victoria: 20mi. NW Swan Hill (Greaves,T.); Bannerton Mallee (Nilson,A.C.); Bendigo (Greenslade,P.J.M.; Lee,K.E.); Ouyen. Comments. This species occurs in semi-arid areas of eastern South Australia, south-central Queensland, western New South Wales and north-western Victoria (Fig. 25). It is generally found in forested habitats including Callitris and mulga (Acacia sp.) woodlands (including on red soils), mallee (with and without Triodia (spinifex grass)) and in savannah woodlands, but is occasionally found in grasslands, especially with scattered trees. It is most often found on sandy soils. Nests are always in soil and almost always have large, deep craters around the entrance. The biology of this species was discussed by Crawley (1922 a: 122).Published as part of Shattuck, Steven O, 2008, Australian ants of the genus Aphaenogaster (Hymenoptera: Formicidae), pp. 25-45 in Zootaxa 1677 on pages 30-31, DOI: 10.5281/zenodo.18032

Aenictus levior Karavaiev, n. stat.

Aenictus levior (Karavaiev), n. stat. Eciton (Aenictus) impressus var. levior Karavaiev, 1926: 425 (junior synonym of A. aratus: Wilson, 1964: 446; new status as full species). Types. Worker syntypes from Buru Island, Indonesia (not examined). Comments. Aenictus material from Indonesia is uncommon in collections and drawing conclusions on the species involved is difficult. However, it would appear that levior is a valid species and not a junior synonym of aratus as suggested earlier. Morphologically this species has a head shape similar to aitkenii (relatively broad) but the scape is shorter. However, this analysis is based on limited material and should be considered tentative until additional specimens become available. Material referable to this species has been examined from Sumatra, Indonesia, peninsular Malaysia and Sarawak, Malaysia.Published as part of Shattuck, Steven O., 2008, Review of the ant genus Aenictus (Hymenoptera: Formicidae) in Australia with notes on A. ceylonicus (Mayr), pp. 1-19 in Zootaxa 1926 on page 17, DOI: 10.5281/zenodo.18481

Aenictus philiporum Wilson

Aenictus philiporum Wilson (Figs 15 –17, 28) Aenictus philiporum Wilson, 1964: 473, fig. 74. Types. Holotype worker and 5 paratype workers from Iron Range, Queensland (MCZC, examined). Diagnosis. Head capsule smooth posteriorly and weakly punctate between the frontal carinae and above the mandibular insertions; pronotum with large smooth areas dorsally and laterally, other areas micro-reticulate. Worker Description. Mandible broad, triangular, with 6–9 widely spaced short teeth; anterior clypeal border broadly convex, even with or slightly posterior to anterior margin of frontal lobes; parafrontal ridges present; subpetiolar process weakly developed, broadly rounded anteriorly, flat posteriorly; head smooth laterally, remainder weakly to moderately punctate; posterior section of pronotum and entire mesonotum completely smooth to very weakly reticulate; anterior of pronotum, mesopleuron and entire propodeum weakly punctuate; body red-brown, head, antennae and legs lighter yellow-red. Measurements. Worker (n = 10) - CI 88–94; HL 0.66–0.71; HW 0.59–0.67; MTL 0.47–0.57; ML 0.96– 1.06; SI 83–89; SL 0.52–0.59. Material examined. Australia: Queensland: Claudie River, Iron Range (Monteith,G.) (ANIC). Papua New Guinea: Morobe: Bulolo (Lowery,B.B.) (ANIC). Comments. This rare species has been collected only a limited number of times on Cape York Peninsula, Queensland and in eastern Papua New Guinea. Nothing is known concerning its biology.Published as part of Shattuck, Steven O., 2008, Review of the ant genus Aenictus (Hymenoptera: Formicidae) in Australia with notes on A. ceylonicus (Mayr), pp. 1-19 in Zootaxa 1926 on page 13, DOI: 10.5281/zenodo.18481

Aenictus hilli Clark

Aenictus hilli Clark Aenictus hilli Clark, 1928: 38. Types. Holotype male from Malanda, Queensland (ANIC, examined). Comments. A. hilli was described by Clark (1928) from a single male collected at Malanda, Queensland. There are numerous males in ANIC which are morphologically similar to the type of A. hilli. Unfortunately none are associated with workers. In addition, the remaining described Australian species are all worker-based and without associated males. This makes it impossible to positively associate A. hilli with any of these other species. Distribution patterns give little clue as to the association either as Aenictus aratus, A. nesiotis, A. prolixus and A. turneri all occur in the general area of the type locality of A. hilli. Thus there is currently insufficient information to associate A. hilli, or any of these other males, with any of the worker-based species. As a result A. hilli is here treated as a valid species and these additional males tentatively associated with it until such time that worker-associated males or fresh material suitable for molecular analysis can be secured. Material examined (all unassociated males). Australia: Northern Territory: 34 mi. NW of Dorisvale HS (Mendum,M.) (ANIC); 39 km E of Alice Springs (Cardale,J.C.) (ANIC); 4 mi. W of Coolibah H.S. (Mendum,M.) (ANIC); 48 mi. SW of Daly River (Mendum,M.) (ANIC); 5mi. ENE Victoria River Downs (Kelsey,L.P.) (ANIC); Daly River Mission (Hutchinson,J.F.) (ANIC); Katherine (Kelsey,L.P.) (ANIC); Todd River, 9 km N by E of Alice Springs (Cardale,J.C.) (ANIC); Queensland: Bamaga (Sedlacek,J.) (ANIC); Lockerbie, Cape York (Sedlacek,J.) (ANIC); Malanda (Hill,G.F.) (ANIC); West Claudie River, Iron Range (Monteith,G.B. & Cook,D.) (ANIC); Western Australia: Carson Escarpment (Common,I.F.B. & Upton,M.S.) (ANIC); Drysdale River (Common,I.F.B. & Upton,M.S.) (ANIC).Published as part of Shattuck, Steven O., 2008, Review of the ant genus Aenictus (Hymenoptera: Formicidae) in Australia with notes on A. ceylonicus (Mayr), pp. 1-19 in Zootaxa 1926 on page 8, DOI: 10.5281/zenodo.18481