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Crematogaster modiglianii Crematogaster modiglianii has a widespread distribution in South-East Asia, but does not occur in Java, while another widespread species, C. seaeardi, occurs also in Java. What could explain the absence of C. modiglianii on Java? Menzel & Blüthgen (2010) revealed parabiotic associations between C. modiglianii and Camponotus (Myrmotarsus) rufifemur Emery, 1900. Camponotus rufifemur is known from the Indochinese Peninsula, Malay Peninsula, Borneo, Sumatra and Philippines, but not distributed in Java (Antmaps, 2021; AntWiki, 2021). The close relationship might have affected the absence of C. modiglianii in Java. Alternately, Java has a different climate from the other islands on the Sunda Shelf (Wilting et al., 2012). The climate of Java is drier than in other parts of the Sunda Shelf (Heaney, 1991). Veron et al. (2007) suggested that open and drier habitat in Java at the time might have affected the species distribution of the mongoose Herpestes species. Land bridges connecting the Sundaic regions might have allowed for the migration of C. modiglianii, but the species did not successfully colonize, probably due to ecological competition with related taxa., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Menzel F, Bluthgen N. 2010. Parabiotic associations between tropical ants: equal partnership or parasitic exploitation? Journal of Animal Ecology 79: 71 - 81.","Antmaps. 2021. Antmaps. Available at: http: // www. antmaps. org / (accessed on 1 March 2021).","Wilting A, Sollmann R, Meijaard E, Helgen KM, Fickel J. 2012. Mentawai's endemic, relictual fauna: is it evidence for Pleistocene extinctions on Sumatra? Journal of Biogeography 39: 1608 - 1620.","Heaney LR. 1991. A synopsis of climatic and vegetational change in South-East-Asia. Climatic Change 19: 53 - 61.","Veron G, Patou M-L, Pothet G, Simberloff D, Jennings AP. 2007. Systematic status and biogeography of the Javan and small Indian mongooses (Herpestidae, Carnovora). Zoologica Scripta 36: 1 - 10."]}

The Crematogaster difformis-subgroup The relationships of the sister-species in the C. difformis -subgroup were variable with respect to their geographical distribution patterns. Crematogaster ampullaris and C. seaeardi display an allopatric distribution, whereas C. difformis and C. tanakai display a sympatric distribution. The closely related C. ampullaris and C. seaeardi illustrate an interesting distribution pattern with C. ampullaris in Sulawesi and C. seaeardi widespread elsewhere in South-East Asia. The estimated dating suggests that C. ampullaris and C. seaeardi diverged approximately 4 Mya (Fig. 4; Table 2). The divergence implies a recent split of these populations and landmass distributions in that geological time (Hall, 2002) suggesting that the ancestor of both species dispersed from the Sundaic region to Sulawesi west-to-east across Wallace’s Line. Sulawesi is considered to have taken its present form due to the collision of different islands c. 15–5 Mya (Lohman et al., 2011). As the last possible connections between Borneo and Sulawesi were during the Pliocene or the Miocene (Morales & Melnick, 1998; Mercer & Roth, 2003), the colonization of Sulawesi by the common ancestor may have occurred much later after the island collision event. The ancestor of C. seaeardi and C. ampullaris may have expanded its range to western Sulawesi (eastern Sunda Shelf) during low sea-level periods and evolved into C. ampullaris. As the Makassar Strait is a deep trench, Borneo and Sulawesi were not connected when sea levels dropped. Their diversification resulted from ‘an inter-island dispersal combined with climatic vicariance event’ (Condamine et al., 2013). Crematogaster difformis and C. tanakai are sympatric in Borneo, and their habitats are spatially close to each other. Crematogaster difformis has an obligate mutualistic relationship with Lecanopteris and Platycerium ferns in canopy trees of lowland rainforests, and C. tanakai inhabits the same nests with C. difformis (Tanaka et al., 2009, 2012; Tanaka & Itioka, 2011). This restricted distribution demonstrates distinct endemism and is, therefore, of conservation significance. Lecanopteris fern species are found in the canopies of lowland tropical rainforests. The preservation of such primary forests will lead to the conservation of the endemic species. Molecular dating indicates that C. difformis and C. tanakai diverged approximately 1 Mya (Fig. 4; Table 2). The low divergence implies a recent speciation. As in C. aurita, the dispersal event and competition can explain similarly the position of C. mucronata, which is endemic to Sumatra (Fig. 4). Klaus et al. (2013) suggested early divergence of freshwater crabs Parathelphusa and subsequent appearance on Sumatra, since much of Sumatra was still submerged during the Late Miocene. In contrast, the ancestor of C. mucronata already appeared on fragmented Sumatra during the Late Miocene. Freshwater crabs require more land area for their habitats, but ants presumably do not require large areas as Yamane (2013) listed 126 ant species having colonized Krakatau Island after eruption. Faunal and genetic similarities between the Malay Peninsula and Sumatran populations have been documented for several taxa (Gorog et al., 2004; Lohman et al., 2011; Leonard et al., 2015), as the two areas were connected at sea level 20–30 m below the present level (Voris, 2000)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Hall R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences 20: 353 - 431.","Lohman DJ, de Bruyn M, Page T, von Rintelen K, Hall R, Ng PKL, Shin H-T, Carvalho GR, von Rintelen T. 2011. Biogeography of the Indo-Australian Archipelago. Annual ReVieae of Ecology, EVolution, and Systematics 42: 205 - 226.","Morales JC, Melnick DJ. 1998. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), as revealed by high resolution restriction site mapping of mitochondrial ribosomal genes. Journal of Human EVolution 34: 1 - 23.","Mercer JM, Roth VL. 2003. The effects of Cenozoic global change on squirrel phylogeny. Science 299: 1568 - 1572.","Condamine FL, Toussaint EFA, Cotton AM, Genson GS, Sperling FAH, Kergoat GJ. 2013. Fine-scale biogeographical and temporal diversification processes of peacock swallowtails (Papilio subgenus Achillides) in the Indo-Australian Archipelago. Cladistics 29: 88 - 111.","Tanaka HO, Inui Y, Itioka T. 2009. Anti-herbivore effects of an ant species, Crematogaster difformis, inhabiting myrmecophytic epiphytes in the canopy of a tropical lowland rainforest in Borneo. Ecological Research 24: 1393 - 1397.","Tanaka HO, Yamane S, Itioka T. 2012. Effects of a ferndwelling ant species, Crematogaster difformis, on the ant assemblages of emergent trees in a Bornean tropical rainforest. Annals of the Entomological Society of America 105: 592 - 598.","Tanaka HO, Itioka T. 2011. Ants inhabiting myrmecophytic ferns regulate the distribution of lianas on emergent trees in a Bornean tropical rainforest. Biology Letters 7: 706 - 709.","Klaus S, Selvandran S, Goh JW, Wowor D, Brandis D, Koller P, Schubart CD, Streit B, Meier R, Ng PKL, Yeo DCJ. 2013. Out of Borneo: Neogene diversification of Sundaic freshwater crabs (Crustacea: Brachyura: Gecarcinucidae: Parathelphusa). Journal of Biogeography 40: 63 - 74.","Yamane SK. 2013. A review of the ant fauna of the Krakatau Islands, Indonesia. Bulletin of the Kitakyushu Museum of Natural History and Human History 11: 1 - 66.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Leonard JA, Den Tex R-J, Hawkins MTR, Munoz-Fuentes V, Thorington R, Maldonado JE. 2015. Phylogeography of vertebrates on the Sunda Shelf: a multi-species comparison. Journal of Biogeography 42: 871 - 879.","Voris HK. 2000. Maps of Pleistocene sea levels in South-East Asia: shorelines, river systems and time durations. Journal of Biogeography 27: 1153 - 1167."]}

The Crematogaster inflata-subgroup Crematogaster aurita is distributed in Indochina, and does not extend southward beyond the Isthmus of Kra. The phylogenetic position of the species was sister to C. onusta within the C. inflata -subgroup. Biogeographic analyses based on the distribution pattern and phylogenetic position of the species suggests that C. aurita appeared as a result of a dispersal event (Fig. 4). During the Pliocene to Pleistocene, terrestrial Indochinese animals might have had many opportunities to disperse to the Sundaic regions and vice versa, since these land areas were connected in the Sunda Shelf during glacial periods. The absence of C. aurita to the south of the Isthmus of Kra presumably resulted from limited dispersal capability and could also be a consequence of competition between related species. The distribution in the Indochinese Peninsula is possibly related to the preference of C. aurita for open, dry habitats. This preference could be explained by savannah corridors that are thought to have spanned Sundaland during glacial periods (Bird et al., 2005) and drier and seasonal habitats are currently present in the Indochinese Peninsula and Java. However, Crematogaster inflata and C. physothorax are sympatric in the Malay Peninsula. Our molecular dating analyses revealed that C. inflata and C. physothorax diverged approximately 6 Mya (Fig. 4; Table 2). This high divergence implies an old split of these populations. It is unclear whether this divergence resulted from a vicariant event or dispersal of the ancestor of both species. The sympatric distribution of these two species in the peninsula might be due to secondary contact after their speciation (Fig. 4). The estimated divergence suggests that the present geographic overlap of the two species is the result of independent colonization events from different geographic centres of origin. Hosoishi & Ogata (2009: figs 25, 28) represented the distribution records of Crematogaster inflata and C. physothorax based on the original descriptions and literature, but the allocations in the figures seem controversial. A record of C. inflata from Myanmar was based on misidentification. Yet extensive field surveys in the Indochinese Peninsula were not able to find C. physothorax (e.g. Khachonpisitsak et al., 2020), whose type locality is Thagata, Tenasserim, Myanmar and presumably this species is not distributed in the Indochinese Peninsula., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Hosoishi S, Ogata K. 2009. A taxonomic revision of the Asian endemic subgenus Physocrema of the genus Crematogaster (Hymenoptera: Formicidae). Zootaxa 2062: 15 - 36.","Khachonpisitsak S, Yamane S, Sriwichai P, Jaitrong W. 2020. An updated checklist of the ants of Thailand (Hymenoptera, Formicidae). ZooKeys 998: 1 - 182."]}

The Crematogaster Ʋacca-subgroup The phylogenetic relationship between the C. Ʋaccasubgroup and their ancestral state reconstruction suggests that their common ancestor occurred in the Sundaic region and its founder dispersed to Sumatra (Fig. 4). During glacial periods, the populations (C. yamanei) retracted into glacial forest refugia. When environmental conditions became favourable for expansion, the populations recolonized, but in the Sundaic region they were restricted to higher elevation areas due to competition with the C. inflata -subgroup and the C. difformis -subgroup, which were already present in this area at lower elevations. Crematogaster yamanei is endemic to Sumatra, but closely related to C. Ʋacca, which is distributed in the Malay Peninsula and Borneo. Meijaard’s (2004) mammal studies suggested that Sumatra shares elements of its fauna with both the Bornean/Javan and Malayan regions as a result of secondary contact. Those patterns are found in the younger taxon C. yamanei, but not in the older taxon C. mucronata. The restricted distributions of C. mucronata and C. yamanei could have resulted from another speciation event and extirpation of the relatives, respectively., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Meijaard E. 2004. SolVing mammalian riddles: a reconstruction of the Tertiary and Quaternary distribution of mammals and their palaeoenVironments in island South-East Asia. PhD Thesis, Australian National University, 349."]}

IS CREMATOGASTER SUBCIRCULARIS A RELICT SPECIES? Crematogaster subcircularis was revealed to have an interesting distribution pattern. This species is found in mountainous areas (600–1250 m elevation) of the Sundaic region: the Malay Peninsula (Fraser’s Hill, 1000 m), Borneo (Poring, 600–700 m; Crocker Range, 1100 m) and Java (Cikaniki, 1000–1250 m). This distribution suggests that the ancestor retreated to high-altitude forests during the ice age, whereas the other related species (C. modiglianii) remained in lowaltitude forests (0–600 m elevation). Despite a postulated savannah corridor through central Sundaland (Bird et al., 2005) that might have restricted the west– east dispersal of forest species, gallery forests in the Sunda Shelf could have served as corridors for forest species and facilitated the dispersal of C. subcircularis (Fig. 4). Rising sea-levels in the Late Pleistocene may have isolated the three populations. Molecular dating estimates and restricted highland distribution imply that C. subcircularis is a relict species. Several studies have hypothesized that continuous mountain ranges, which span Myanmar, Thailand, Laos, south China and Vietnam, acted as glacial refugia (Brandon-Jones, 1996; Gorog et al., 2004; Iyengar et al., 2005). Assuming that C. subcircularis is a relict with a wide distribution range in the glacial periods, those populations were fragmented into several geographical ones when the sea level was higher, then closely related or sister-species might have retreated to the high-elevation refugia of mainland South-East Asia (Surridge et al., 1999)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on pages 917-918, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Brandon-Jones D. 1996. The Asian Colobinae (Mammalia, Cercopithecidae) as indicators of Quaternary climatic changes. Biological Journal of the Linnean Society 59: 327 - 350.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Iyengar A, Babu N, Hedges S, Venkataraman A, Maclean N, Morin PA. 2005. Phylogeography, genetic structure and diversity in the dhole (Cuon alpinus). Molecular Ecology 14: 2281 - 2297.","Surridge AK, Timmins RJ, Hewitt GM, Bell DJ. 1999. Striped rabbits in South-East Asia. Nature 400: 726 - 726."]}

This is the first discovery of nesting an ant queen in the mantid’s oothecae chamber. A total of 93 oothecae have been collected and dissected. After tearing the outer skin of the most oothecae apart (80 out of 93), a queen ant Crematogaster inermis (Formicidae: Crematogastrini) and her eggs were observed. All of these 80 oothecae were parasited by parasitoid wasp Podagrion pachymerum (Chalcidoidea: Toryminae). According to our observations, it was assumed that the ant, Crematogaster inermis (Formicidae: Crematogastrini), inhabited the oothecae of mantid, Hierodula tenuidentata (Mantidae: Paramantini), to get a ready nest and accessible food using parasitoid wasps. We observed a mixed-species aggregation of arthropods in most of the examined oothecae. The arthropods that were observed in most of the collected oothecae are listed below: An ant, Crematogaster inermis (Formicidae: Crematogastrini); a mimicking spider ant, Synageles persianus (Salticidae: Dendryphantini); a parasitoid, Podagrion pachymerum (Chalcidoidea: Toryminae); a pseudoscorpion species, Chernes sp. (Chernetidae: Chernetini); a thysanoptera species, Haplothrips sp. (Phlaeothripidae: Haplothripini) and a neuroptera species, Semidalis sp. (Coniopterygidae: Conwentziini).

Crematogaster jacindae Sharaf & Hita Garcia, 2019 Fig. 32 Crematogaster jacindae Sharaf & Hita Garcia, 2019 in Sharaf et al. 2019: 58 (w) Oman. Afrotropic. Diagnosis Bicolored species, head black-brown to black, mesosoma, petiole, and postpetiole dark brown, gaster golden yellow; area in front of eyes finely longitudinally striated; cephalic surface feebly imbricate; eyes with about 11 ommatidia in longest row; mesonotum without tubercle; mesopleura and metapleura clearly imbricate; mesonotum with a single pair of setae; propodeal spiracles slit-shaped. Material examined OMAN – Dhofar • w, holotype; Ayn Sahlanot; 17.148° N, 54.179° E; alt. 151 m; 16 Nov. 2017; M.R. Sharaf leg.; KSMA • w, paratype; Ayn Hamran; 17.086° N, 54.280° E; alt. 56 m; 22 Nov. 2017; BS; M.R. Sharaf leg.; KSMA • 11 w; same collection data as for paratype; KSMA • 1 w, paratype; same collection data as for paratype; CASENT0922856; CASC • 1 w; same collection data as for paratype; WMLC • 8 w; Ayn Razat; 17.124° N, 54.238° E; alt. 98 m; 20 Nov. 2017; ML; M.R. Sharaf leg.; KSMA • 16 w; Dhalkout; 16.727° N, 53.249° E; alt. 623 m; 18 Nov. 2017; SF; M.R. Sharaf leg.; KSMA • 7 w; Dhalkout; 16.707° N, 53.251° E; alt. 34 m; 19 Nov. 2017; BS; M.R. Sharaf leg.; KSMA • 17 w; Dhalkout; 16.693° N, 53.156° E; alt. 628 m; 18 Nov. 2017; BS; M.R. Sharaf leg.; KSMA • 4 w; Ayn Sahlanot; 17.148° N, 54.179° E; alt. 151 m; 16 Nov. 2017; BS; M.R. Sharaf leg.; KSMA • 20 w; Ayn Ashat; 16.998° N, 53.820° E; alt. 202 m; 21 Nov. 2017; SF; M.R. Sharaf leg.; KSMA • 1 w; Dhalkout Road, Aghbaroot Village; 16.798° N, 53.554° E; alt. 1034 m; 18 Nov. 2017; BS; M.R. Sharaf leg.; KSMA. Ecological and biological notes The microhabitats of C. jacindae included leaf litter, soil, under stones, or on native vegetation, especially Vachellia. The majority of specimens were collected foraging on plants using a beating sheet. Workers were observed foraging on ground and wild shrubs (Sharaf et al. 2019). Geographic range Only known from Oman., Published as part of Sharaf, Mostafa R., Wetterer, James K., Mohamed, Amr A. & Aldawood, Abdulrahman S., 2022, Faunal composition, diversity, and distribution of ants (Hymenoptera: Formicidae) of Dhofar Governorate, Oman, with updated list of the Omani species and remarks on zoogeography, pp. 1-106 in European Journal of Taxonomy 838 (1) on page 52, DOI: 10.5852/ejt.2022.838.1925, http://zenodo.org/record/7094306, {"references":["Sharaf M. R., Aldawood A. S. & Hita Garcia F. 2019. Review of the Arabian Crematogaster Lund (Hymenoptera, Formicidae), synoptic list, distribution, and description of two new species from Oman and Saudi Arabia. ZooKeys 898: 27 - 81. https: // doi. org / 10.3897 / zookeys. 898.37531"]}

Crematogaster chiarinii Emery, 1881 Fig. 31 Crematogaster chiarinii Emery, 1881a: 271 (w) Ethiopia. Afrotropic. Diagnosis Head, petiole, postpetiole, and gaster dark brown, mesosoma light brown; head, in full-face view, with a well-defined frontal triangle and posterior longitudinal carina reaching posterior margin of eyes; propodeal dorsum transversally striated in dorsal view; postpetiole seen from above as broad anteriorly as posteriorly. Material examined This species was not collected during the present study. Ecological and biological notes This species forms large colonies with workers foraging among myrmecophilous myrmedoniine beetles at the base of trees of Vachellia (Collingwood 1985). Geographic range Crematogaster chiarinii was originally described from Ethiopia and elsewhere in north-east Africa. It was later recorded from Oman, KSA, and Yemen (Collingwood 1985; Collingwood & Agosti 1996; Collingwood & van Harten 2001). This species has been recorded from Dhofar by Collingwood (1985) based on a single specimen., Published as part of Sharaf, Mostafa R., Wetterer, James K., Mohamed, Amr A. & Aldawood, Abdulrahman S., 2022, Faunal composition, diversity, and distribution of ants (Hymenoptera: Formicidae) of Dhofar Governorate, Oman, with updated list of the Omani species and remarks on zoogeography, pp. 1-106 in European Journal of Taxonomy 838 (1) on page 49, DOI: 10.5852/ejt.2022.838.1925, http://zenodo.org/record/7094306, {"references":["Emery C. 1881 a. Spedizione italiana nell'Africa equatoriale. Risultati zoologici. Formiche. Annali del Museo Civico di Storia Naturale Giacomo Doria 16: 270 - 276.","Collingwood C. A. 1985. Hymenoptera: Fam. Formicidae of Saudi Arabia. Fauna of Saudi Arabia 7: 230 - 302.","Collingwood C. A. & Agosti D. 1996. Formicidae (Insecta: Hymenoptera) of Saudi Arabia (part 2). Fauna of Saudi Arabia 15: 300 - 385.","Collingwood C. A. & van Harten A. 2001. Additions to the ant fauna of Yemen (Hymenoptera, Formicidae). Esperiana. Buchreihe zur Entomologie 8: 559 - 568."]}

Crematogaster acaciae Forel, 1892 Fig. 30 Crematogaster acaciae Forel, 1892: 141 (w) Ethiopia. Afrotropic. Diagnosis Body uniform yellow; head, in full-face view, with a shallow frontal triangle and without longitudinal carina; postpetiole in dorsal view broader posteriorly than anteriorly; propodeal dorsum seen from above longitudinally striated. Material examined This species was not collected during the present study. Ecological and biological notes Nothing has been reported on the ecology of this species. Geographic range Originally described from Ethiopia and subsequently recorded from Oman (Collingwood 1985; Collingwood & Agosti 1996). This species was previously recorded from Dhofar based on two workers (Collingwood 1985)., Published as part of Sharaf, Mostafa R., Wetterer, James K., Mohamed, Amr A. & Aldawood, Abdulrahman S., 2022, Faunal composition, diversity, and distribution of ants (Hymenoptera: Formicidae) of Dhofar Governorate, Oman, with updated list of the Omani species and remarks on zoogeography, pp. 1-106 in European Journal of Taxonomy 838 (1) on page 49, DOI: 10.5852/ejt.2022.838.1925, http://zenodo.org/record/7094306, {"references":["Forel A. 1892. Die Akazien- Crematogaster von Prof. Keller aus dem Somaliland. Zoologischer Anzeiger 15: 140 - 143.","Collingwood C. A. 1985. Hymenoptera: Fam. Formicidae of Saudi Arabia. Fauna of Saudi Arabia 7: 230 - 302.","Collingwood C. A. & Agosti D. 1996. Formicidae (Insecta: Hymenoptera) of Saudi Arabia (part 2). Fauna of Saudi Arabia 15: 300 - 385."]}

CREMATOGASTER OPUNTIAE BUREN, 1968, STAT. REV. (FIG. 25) Crematogaster opuntiae Buren, 1968: 120. Holotype worker, Santa Rita Experimental Range, Arizona (Buren) (LACM) (examined). Junior synonym of C. vermiculata: Morgan & Mackay, 2017: 396; here overturned. Worker measurements ( N = 10): HW 0.86–1.21, HL 0.80–1.11, SL 0.72–0.94, WL 0.89–1.27, MtFL 0.74– 1.05, MSC 1–3, A4SC 5–16, PP-SL/HW 0.14–0.19, CI 1.05–1.11, OI 0.26–0.29, SI 0.75–0.84, MtFL/HW 0.85– 0.89, SPL/HW 0.19–0.25, SPTD/HW 0.50–0.59. Discussion: Crematogaster opuntiae bears little similarity or close phylogenetic relationship (Fig. 1) to the eastern, swamp-inhabiting species C. vermiculata, under which it was synonymized by Morgan & Mackay (2017). Among obvious differences, it has longer scapes and legs (SL/HL 0.83–0.90 and MtFL/HL 0.91–0.95 compared to SL/HL 0.75–0.82 and MtFL/HL 0.84–0.89 in C. vermiculata), predominantly reticulate-foveolate sculpture on the promesonotum, more divergent propodeal spines (SPTD/ PPW 1.71–1.96 compared to 1.15–1.48 in C. vermiculata) and lesser amounts of standing pilosity (MSC 1–3 and A4SC 5–16, compared to MSC 3–9 and A4SC 9–20 in C. vermiculata). As noted by Buren (1968), C. opuntiae is similar to C. californica, differing primarily by the shinier head and by the more appressed pubescence on the scapes and head. The relationship of C. opuntiae to C. californica remains to be clarified – they may well prove to be conspecific. Crematogaster opuntiae has similar eye, scape, leg (Fig. 42) and propodeal spine dimensions as C. californica, and in our phylogenomic analyses the two are sister-taxa (Fig. 1). Distribution and biology: Buren (1968) recorded C. opuntiae from desert and semi-desert regions of Arizona and considered C. californica to be restricted to California and Baja California. Both taxa are ground-nesting., Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on page 919, DOI: 10.1093/zoolinnean/zlab047, {"references":["Buren WF. 1968. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia Entomological Society 3: 91 - 121.","Morgan C, Mackay W. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Balti, Moldova: Lambert Academic Publishing."]}

CREMATOGASTER OPACA MAYR, 1870 Crematogaster opaca Mayr, 1870: 992. Syntype worker, Tabukaya, Mexico, 1865 (Bilimek) (NHMW) (examined via AntWeb image: CASENT0919705) and syntype worker, Mexico (Sichel) (BMNH) (examined via AntWeb image: CASENT0902152); latter here designated lectotype. Crematogaster lineolata subsp. opaca Mayr; Emery, 1895: 283. Crematogaster opaca Mayr; Wheeler, 1919: 111. Crematogaster (Acrocoelia) lineolata subsp. opaca Mayr; Emery, 1922: 141. Crematogaster (Acrocoelia) opaca Mayr; Wheeler, 1934a: 139. Crematogaster (Crematogaster) opaca Mayr; Smith, 1958: 126. Discussion: Morgan & Mackay (2017) refer to a syntype worker of C. opaca in NHMW ‘collected by Professor Bilimek and Dr Sichel in México’. This statement is misleading because there is no single syntype collected by Bilimek and Sichel; their collections were separate. In the BMNH there is a syntype worker (here designated lectotype) (CASENT0902152) labelled ‘Mex.|Sichel’, while the NHMW syntype (now paralectotype) is labelled ‘ Mexico |Takubaya|Bilim. 65’. However, the important point is that these two specimens are almost certainly not conspecific. The Sichel specimen has the head, mesosoma, petiole and postpetiole densely reticulate-foveolate and opaque, and the gaster finely reticulate and sublucid; the promesonotum is evenly convex in profile and broad in dorsal view (pronotum width/head width ~0.62); the propodeal spine distance is relatively large (SPTD/HW ~0.49); and standing pilosity appears to be sparse (not visibly present on the mesosoma dorsum). The Bilimek specimen is more weakly sculptured with a correspondingly shinier integument, especially on the petiole, postpetiole and gaster; the promesonotum is somewhat flattened in profile and narrower in dorsal view (pronotum width/ head width ~0.53); the propodeal spine tips are less distant (SPTD/HW ~0.44); and standing pilosity is more conspicuous, being present on the mesosoma dorsum on both the pronotal humeri and the mesonotal declivity. Although in this study we are concerned primarily with the Crematogaster of America north of Mexico, we have decided to select a lectotype of C. opaca to assist future work. We designate the BMNH specimen as lectotype because it better matches the original description, which emphasizes the densely reticulate-punctate sculpture and the scarcity of standing pilosity (Mayr, 1870). Unfortunately the provenance of this specimen – other than Mexico – is unknown. Sichel evidently supplied the specimen to Mayr, but there is no evidence that he visited Mexico or anywhere else outside Europe during his lifetime (Mulsant, 1869), so the specimen would have been acquired from another collector, possibly de Saussure. The non-conspecific paralectotype is from Takubaya (= Tacubaya), a district of Mexico City, near Bilimek’s place of residence during his stay in Mexico (Roth, 1965: 346). There is a confusing array of C. opaca -like forms found in Mexico and Central America, showing considerable variation in sculpture, pilosity, configuration of the promesonotum, size of propodeal spines and petiole shape. More comprehensive sampling and careful study is necessary to determine how many species are present in this complex. These ants just barely enter the United States, in southern Arizona. Because of taxonomic uncertainty, we refer to the Arizona populations as Crematogaster sp. cf. opaca. These specimens are similar to the C. opaca lectotype with respect to integument sculpture, promesonotal shape (pronotum width/head width 0.59–0.62) and propodeal spine distance (SPTD/ HW 0.45–0.51), but they have more abundant standing pilosity (MSC 2–5, A4SC 10–18)., Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on pages 918-919, DOI: 10.1093/zoolinnean/zlab047, {"references":["Mayr G. 1870. Neue Formiciden. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 20: 939 - 996.","Wheeler WM. 1919. A new paper-making Crematogaster from the south-eastern United States. Psyche (Cambridge) 26: 107 - 114.","Emery C. 1922. Hymenoptera, fam. Formicidae, subfam. Myrmicinae. Genera Insectorum 174 B: 95 - 206.","Wheeler WM. 1934 a. Ants from the islands off the west coast of Lower California and Mexico. Pan-Pacific Entomologist 10: 132 - 144.","Smith MR. 1958. Family Formicidae. Pp. 108 - 162 in: Krombein KV, ed. Hymenoptera of America north of Mexico. Synoptic catalogue. First supplement. United States Department of Agriculture. Agriculture Monograph 2 (suppl. 1): 1 - 305.","Morgan C, Mackay W. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Balti, Moldova: Lambert Academic Publishing.","Mulsant E. 1869. Notice sur le Dr. Jules Sichel. Annales de la Societe Linneenne de Lyon 17: 383 - 410.","Roth PHJ. 1965. Dominik Bilimek Leben und Werk eines osterreichischen Naturforschers: Zur mexikanischen Expedition der Osterreicher vor hundert Jahren. Sudhoffs Archiv fur Geschichte der Medizin und der Naturwissenschaften 49: 338 - 354."]}

CREMATOGASTER CERASI (FITCH, 1855) (FIGS 10B, 31) Myrmica cerasi Fitch, 1855: 835. Four syntype workers, New York (USNM) (examined); one worker here designated lectotype (USNMENT00529078) (image on AntWeb). Crematogaster cerasi (Fitch, 1855); Roger, 1863: 37. Junior synonym of Crematogaster lineolata (Say): Dalla Torre, 1893: 83; Creighton 1950: 213. Crematogaster lineolata var. cerasi (Fitch): Emery, 1895: 282. Crematogaster lineolata cerasi (Fitch): Smith, 1951: 809. Status as species: Buren in Smith, 1958: 125; Buren, 1968: 92; Johnson, 1988: 318. Senior synonym of Crematogaster kennedyi: Buren in Smith, 1958: 125. Senior synonym of Crematogaster browni: Morgan & Mackay, 2017: 91; here overturned (see above). Worker measurements ( N = 17): HW 0.77–1.05, HL 0.74–0.99, SL 0.62–0.82, WL 0.83–1.17, MtFL 0.67– 0.95, MSC 2–7, A4SC 4–14, PP-SL/HW 0.15–0.19, CI 1.03–1.13, OI 0.22–0.25, SI 0.72–0.81, MtFL/HW 0.81– 0.92, SPL/HW 0.22–0.25, SPTD/HW 0.45–0.59. Discussion: This is a common species in eastern North America that overlaps broadly in distribution with C. lineolata. It is distinguished from the latter by having the standing pilosity on the mesosoma dorsum restricted to one to several setae on each pronotal humerus (MSC 2–7 vs. 15–26 in C. lineolata). The pilosity on the gaster is also sparser in C. cerasi (A4SC 4–14 vs. 16–42 in C. lineolata) but the postpetiolar seta is longer (PP-SL/PPW 0.42–0.54 vs. 0.28–0.43 in C. lineolata). For differences between C. cerasi and C. browni, see under the latter species. Type notes: The four syntypes of C. cerasi in USNM are old and in poor condition; each is on a separate pin and labeled ‘N. Y.’, ‘Collection| T Pergande’ and ‘No 53583| U.S. N.M. |Cotype’. The specimens have also been assigned recent USNMENT specimen codes: 00921679, 00921680, 00921681 and 00529078. The last is in best condition and is here designated lectotype. Specific measurements of the lectotype are as follows: HW 1.05, HL 0.94, SL 0.80, ED 0.26, WL 1.14, MtFL 0.92, SPL 0.26, SPTD 0.61. Distribution and biology: Crematogaster cerasi is widespread in eastern North America, from southern Canada to Florida. It occurs as far west as North Dakota (Wheeler & Wheeler, 1977), Colorado (Gregg, 1963; as ‘ Crematogaster lineolata ’) and northern New Mexico (material examined in LACM). Records on AntWeb of ‘ C. lineolata’ from Montana (FMNHINS0000108303) and South Dakota (FMNHINS0000109880) almost certainly refer to C. cerasi. This species nests in rotten logs, in stumps, under stones and directly in the soil, in both open habitats (grasslands, fields) and in woodlands (Wheeler & Wheeler, 1963; Johnson, 1988; Ellison et al., 2012; Deyrup, 2017). It is also known to colonize wooden structures in houses (Smith, 1965)., Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on page 912, DOI: 10.1093/zoolinnean/zlab047, {"references":["Fitch A. 1855. Report on the noxious, beneficial and other insects of the state of New-York. Albany: C. van Benthuysen.","Roger J. 1863. Verzeichniss der Formiciden-Gattungen und Arten. Berliner Entomologische Zeitschrift 7: 1 - 65.","Dalla Torre KW. 1893. Catalogus hymenopterorum hucusque descriptorum systematicus et synonymicus, Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann.","Creighton WS. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology 104: 1 - 585.","Smith MR. 1951. Family Formicidae. Pp. 778 - 875 in: Muesebeck CF, Krombein KV, Townes HK, eds. Hymenoptera of America north of Mexico. Synoptic catalogue. United States Department of Agriculture. Agriculture Monograph 2: 1 - 1420.","Smith MR. 1958. Family Formicidae. Pp. 108 - 162 in: Krombein KV, ed. Hymenoptera of America north of Mexico. Synoptic catalogue. First supplement. United States Department of Agriculture. Agriculture Monograph 2 (suppl. 1): 1 - 305.","Buren WF. 1968. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia Entomological Society 3: 91 - 121.","Johnson C. 1988. Species identification in the eastern Crematogaster (Hymenoptera: Formicidae). Journal of Entomological Science 23: 314 - 332.","Morgan C, Mackay W. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Balti, Moldova: Lambert Academic Publishing.","Wheeler GC, Wheeler J. 1977. North Dakota ants updated. Reno: published by the authors.","Gregg RE. 1963. The ants of Colorado; with reference to their ecology, taxonomy, and geographic distribution. Boulder: University of Colorado Press.","Wheeler GC, Wheeler J. 1963. The ants of North Dakota. Grand Forks: University of North Dakota Press.","Ellison AM, Gotelli NJ, Farnsworth EJ, Alpert GD. 2012. A field guide to the ants of New England. New Haven: Yale University Press.","Deyrup M. 2017. Ants of Florida: identification and natural history. Boca Raton: CRC Press.","Smith MR. 1965. House-infesting ants of the eastern United States. Their recognition, biology, and economic importance. United States Department of Agriculture, Technical Bulletin 1326: 1 - 105."]}

CREMATOGASTER BROWNI BUREN, 1968, STAT. REV. (FIGS 10A, 29), Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on page 907, DOI: 10.1093/zoolinnean/zlab047, {"references":["Buren WF. 1968. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia Entomological Society 3: 91 - 121."]}

CREMATOGASTER LARREAE BUREN, 1968, STAT. REV. (FIG. 22) Crematogaster larreae Buren, 1968: 117. Holotype male, Franklin Mts., El Paso, Texas (W. F. Buren) (LACM) (examined). Junior synonym of C. depilis: Morgan & Mackay, 2017: 155; here overturned. Worker measurements ( N = 5): HW 0.74–1.05, HL 0.71– 0.99, SL 0.59–0.78, WL 0.80–1.09, MtFL 0.63–0.87, MSC 0, A4SC 0, PP-SL/HW 0.10–0.13, CI 1.00–1.06, OI 0.27–0.31, SI 0.74–0.80, MtFL/HW 0.83–0.85, SPL/ HW 0.19–0.23, SPTD/HW 0.48–0.53. Discussion: Crematogaster larreae was described by Buren (1968) as a ‘cryptic’ species closely related to and sympatric with C. depilis, having distinctive queens and males but workers that are very similar to those of C. depilis. The primary difference between the workers of the two species is their colour: C. larreae workers have the head and mesosoma dark brown and the gaster brownish-black, while in C. depilis the head and mesosoma vary from yellowish-brown to reddishbrown (the head may be darker than the mesosoma), and the gaster is medium to dark brown. However, this distinction is tenous and difficult to ascertain in older, faded specimens. Buren (1968) also provided evidence of differences in biology, with C. larreae apparently confined to nesting at the base of creosote bush (Larrea tridentata), in contrast to the generalized ground-nesting habits of C. depilis. Because they could not distinguish the workers of these two forms, Mackay & Morgan (2017) synonymized C. larreae under C. depilis, while making no comment about the male and queen differences. We have sequenced sympatric samples taken in Chihuahuan desert near Portal, Arizona, corresponding to the C. depilis colour phenotype (PSW14684) and the C. larreae colour phenotype (PSW15960 and PSW16636), and we find that they are positioned in different parts of the UCE phylogeny (Fig. 1). The C. larreae -like samples are sister to C. mutans, while the C. depilis - like sample clusters with other C. depilis populations from New Mexico, Arizona and California. Thus we have provisional evidence that there are indeed two species, with C. larreae actually being more closely related to C. mutans (with which it is allopatric) than to C. depilis. On the basis of univariate observations and bivariate plots we have been unable to discover diagnostic morphometric differences between the workers of C. depilis and C. larreae, but it is possible that a more detailed, multivariate analysis would permit discrimination of the workers of these two species. This was found to be effective, for example, to separate two closely similar sister-species in Europe, C. scutellaris and C. schmidti (Mayr, 1853) that also differ subtly in colour (Seifert, 2018). Distribution and biology: Crematogaster larreae was recorded by Buren (1968) from desert localities in west Texas, Arizona and California, nesting in the lower stems and roots of Larrea divaricata. The one nest series that we have collected (PSW15960) was in a similar situation in south-eastern Arizona. Buren (1968) further noted that C. larreae was often absent from localities with Larrea, and he suggested that competition with other desert-dwelling ants may restrict it to harsher, rockier sites. It is also interesting to note that the queens and males of C. larreae are smaller and shinier than those of C. depilis, suggesting perhaps an unusual mode of colony foundation., Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on pages 917-918, DOI: 10.1093/zoolinnean/zlab047, {"references":["Buren WF. 1968. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia Entomological Society 3: 91 - 121.","Morgan C, Mackay W. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Balti, Moldova: Lambert Academic Publishing.","Seifert B. 2018. The ants of central and north Europe. Tauer: Lutra Verlags- und Vertriebsgesellschaft."]}

CREMATOGASTER ROSSI BUREN, 1968, STAT. REV. Crematogaster rossi Buren, 1968: 111. Holotype worker, San Jose del Cabo, Baja California Sur, Mexico (USNM) (USNMENT00531965) (examined). Junior synonym of C. vermiculata: Morgan & Mackay, 2017: 396; here overturned. Worker measurements ( N = 6): HW 0.90–1.1, HL 0.83– 1.04, SL 0.76–0.86, WL 0.95–1.25, MtFL 0.80–0.99, MSC 0–8, A4SC 25–34, PP-SL/HW 0.15–0.19, CI 1.06– 1.13, OI 0.25–0.27, SI 0.78–0.85, MtFL/HW 0.87–0.92, SPL/HW 0.24–0.26, SPTD/HW 0.53–0.60. Discussion: Phylogenomic (UCE) data reveal a close relationship of C. rossi to C.californica and C. opuntiae; it has no close affinity to C. vermiculata (Fig. 1). Crematogaster rossi is recognizable by the combination of a predominantly shiny head, conspicuously rugulose promesonotum on a reticulate-foveolate background and abundant standing pilosity on the gaster. The mesosomal sculpture is superficially similar to that of C. vermiculata, but the two species differ in numerous ways. Crematogaster rossi is larger (HW 0.90–1.10, WL 0.95–1.25), with disproportionately longer scapes (SI 0.78–0.85) and legs (MtFL/HW 0.87–0.92) and with propodeal spines that are straight, longer and more divergent posteriorly (SPL/WL 0.22–0.24, SPTD/HW 0.53–0.60) (compare with equivalent measurements for C. vermiculata below). There are up to two isolated standing hairs on each pronotal humerus and usually a few short standing hairs on the mesonotal declivity (MSC 0–7), but the pilosity on the gaster is particularly well-developed (A4SC 25–34, compared with A4SC 9–20 in C. vermiculata). From the closely related species, C. californica and C. opuntiae, C. rossi can be distinguished primarily by its promesonotal sculpture and abundant standing pilosity on the gaster. Distribution and biology: Crematogaster rossi is known from the Mexican states of Baja California Sur and Baja California, and (more doubtfully) Nayarit. Specimens with habitat information come from ‘oak woodland’ and ‘dry wash next to oak woodland’ but others have been collected from sites in more xeric locations. The species is apparently a ground-nester., Published as part of Ward, Philip S. & Blaimer, Bonnie B., 2022, Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group (Formicidae: Hymenoptera), pp. 893-937 in Zoological Journal of the Linnean Society 194 on page 926, DOI: 10.1093/zoolinnean/zlab047, {"references":["Buren WF. 1968. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia Entomological Society 3: 91 - 121.","Morgan C, Mackay W. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Balti, Moldova: Lambert Academic Publishing."]}