Crematogaster (subgenus)

A nominal subgenus of Crematogaster. The group was last evaluated by Blaimer (2012). Her research and numerous published studies have greatly advanced our understanding of Crematogaster systematics. Blaimer proposed the genus should be divided into two subgenera, Crematogaster and Orthocrema. This rid the genus of thirteen subgenera that were part of an archaic classification system with numerous polyphyletic clades. The two subgenera still represent an incomplete understanding of the relationships between the species of this diverse genus. For example, molecular evidence showed that the subgenus Crematogaster can be divided into two clades. Blaimer suggested these two distinctive groups be referred to as the "global clade” and “Australo-Asian clade.”

Blaimer (2012) stated: The new two-subgenera system as proposed follows a deep molecular divergence event between Orthocrema and Crematogaster sensu stricto, tracing back into the Mid-Eocene (ca.40–45 Mya) when these two clades last shared a common ancestor (Blaimer, in press). Recognizing these two groups from each other on a morphological basis is fairly easily achieved, although a diagnosis of the Crematogaster sensu stricto is more problematic given the high morphological variability. An obvious further improvement to the system presented here would have been the recognition of the global Crematogaster clade and the Australo-Asian Crematogaster clade as separate subgeneric entities, as they certainly show enough molecular divergence to justify a three-subgenera-approach. I refrained from such a formal distinction since it was not possible to assign species to these groups on a morphological basis. I primarily aimed to develop a practical, ‘user-friendly’ classification. Petiole and postpetiolar characters that are useful for the distinction of Crematogaster sensu stricto and Orthocrema become hypervariable within the two sensu stricto clades and are not informative at deeper phylogenetic levels. Similar character patterns, for example the presence/absence of a median longitudinal impression on the postpetiole appear to have evolved multiple times within the genus, but the selective pressures acting on morphological evolution in Crematogaster have yet to be revealed. More biological data will need to be collected to investigate these questions.

The proposed species-groups are intended as a guideline on how to structure the large Crematogaster sensu stricto clade more finely, given the phylogenetic framework. I did not attempt to assign every species sampled within the phylogeny (or otherwise known to me) to a species-group. Many more natural groups will only become apparent when the phylogeny of the genus is extended and should thus be established sequentially as data become available. For very similar reasons, I have not yet proposed species-groups for Orthocrema. The molecular sampling for Orthocrema in Blaimer (in press) is too sparse to make predictions on species-groups, as is obvious from the overall long phylogenetic branches that separate species in this clade from each other. Finally, I conclude that the main advantage of recognizing monophyletic species-groups as suggested here is, that these present manageable subunits for taxonomic revisions. In addition, these species-groups will pose excellent subjects for (needed) comparative ecological and behavioural studies within Crematogaster.

Crematogaster ranavalonae group
Crematogaster ranavalonae group

Key to Asian Crematogaster ranavalonae group species

Crematogaster castanea group
Afrotropical Crematogaster castanea group

Includes the Madagascar Crematogaster degeeri species-assemblage.

Crematogaster hova group
Key to Crematogaster hova-group of Madagascar

Includes members of previous subgenus Decacrema from Africa and Madagascar, in addition to newly described as follows. Crematogaster hova-complex (C. hova Forel, C. schencki Forel, C. ensifera Forel), C. sisa Blaimer, C. mahery Blaimer, C. malala Blaimer, C. grevei Forel, C. sabatra Blaimer, C. nosibeensis Forel and C. liengmei Forel.

This group contains species that were once part of the subgenus Decacrema. That subgenus is no longer valid and its species are no longer organized under a single species-group. The Madagascar Decacrema were placed in the hova group by Blaimer (2012c).

1. Antennae 10-segmented (vs. 11-segments in most other species).

2. Petiole broadly or moderately flared, suboval or hexagonal (Figs 15a, 14a, 18a, 22a, respectively).

3. Postpetiole bilobed, with a more or less distinct median impression (e.g. Figs 14b &15b).

Molecular and morphological data strongly support the monophyly of this group. These ants are found throughout all forest habitats within the five biomes on the island. Nesting habits are arboreal except for one species that is known to occasionally nest on the ground and is adapted to drier habitats. Many of the canopy-nesting species construct carton nests around small branches or attached to tree trunks, or alternatively nest in dead branches or twigs. It is unclear whether these nesting habits are species-specific. Malagasy hova-group species are medium to large-sized ants compared to other Crematogaster species in this region, and they often seem to be the dominant element of the Crematogaster, or even of the entire canopy ant fauna where they occur. Partly this may be mediated by the carton-nesting ability that allows for large and often polydomous colonies. Carton nesters often tend and house mealybugs in the main nest or in special carton shelters, and both twig- and carton-nesting species can further be found living in association with myrmecophilous beetles (Blaimer, pers. observ.).

Crematogaster irritabilis group
1. Petiole hexagonal, suboval, or elongate-rectangular.

2. Postpetiole varying from narrowly bilobed with sharp but narrow median impression, to strongly bilobed with broad median impression.

3. Pronotum laterally rounded or subangular.

4. Mesonotum in lateral view rounded, and without distinct posterior face.

5. Fairly abundant erect pilosity over entire body.

6. Propodeal spines, if present, often with a downward curve.


 * Crematogaster irritabilis
 * Crematogaster polita
 * Crematogaster obnigra
 * ''Crematogaster treubi
 * + several unidentified (or undescribed) species.

Crematogaster kelleri group
Crematogaster kelleri group

Crematogaster inflata group
1. Members either with propodeum greatly inflated, or posterolateral denticles on the mesonotum. 2. Head large and broad, often medially depressed. 3. Petiole elongate-rectangular, oval or circular. 4. Postpetiole of globular shape, a faint posterior impression may be present.

This group most likely includes members of the former subgenera Physocrema and Paracrema (as listed by Bolton, 2011), as well as probably a number of species that have previously been associated with the subgenus Crematogaster sensu stricto.

Definitive membership of species is difficult to assess at this point and only certain for the following:


 * Crematogaster subcircularis
 * Crematogaster inflata
 * Crematogaster onusta
 * Crematogaster ampullaris
 * Crematogaster modiglianii

Crematogaster borneensis group
1. Antennae 10-segmented. 2. Head shape rectangular, longer than wide. 3. Petiole in dorsal view suboval (Fig. 18a), circular (Fig. 21a) or hexagonal (Fig. 22a). 4. Postpetiole globular (e.g. Figs 21b, 24, 25), a faint posterior impression may be present.

Most members of this group are obligate inhabitants of Macaranga plants.


 * Crematogaster borneensis
 * Crematogaster decamera
 * Crematogaster captiosa
 * + several undescribed species

Crematogaster rothneyi group
The Crematogaster rothneyi group can be distinguished from other Asian Crematogaster species by the following characters: (1) mandible with four teeth; (2) 3-segmented antennal club; (3) a deep metanotal groove; (4) developed propodeal spines; (5) petiole broader anteriorly; (6) bilobed postpetiole with distinct longitudinal median sulcus; (7) punctuated mesosoma; (8) stout body setae. This species group is similar to Crematogaster coriaria in having features (3), (4), (5) and (7), but can be distinguished from Crematogaster coriaria by having features (1), (2), (6) and (8) (Hosoishi and Ogata 2015).


 * Crematogaster rothneyi
 * Crematogaster rothneyi haputalensis
 * Crematogaster yaharai

Crematogaster tetracantha group
1. Pronotum antero-laterally with distinct sharp margins or denticles or spinose protuberances. 2. Petiole in dorsal view either greatly flared and transversely convex, or suboval.

This group comprises all members of the previous subgenus Xiphocrema and some additional species from Crematogaster.


 * Crematogaster dahlii
 * Crematogaster tetracantha
 * Crematogaster recurva
 * Crematogaster elegans
 * Crematogaster flavitarsis
 * Crematogaster weberi
 * Crematogaster elysii
 * Crematogaster abrupta

Nomenclature

 *  CREMATOGASTER [Myrmicinae: Crematogastrini]
 * Crematogaster Lund, 1831a: 132. Type-species: Formica scutellaris, by subsequent designation of Bingham, 1903: 124.
 * ACROCOELIA [junior synonym of Crematogaster (Crematogaster)]
 * Acrocoelia Mayr, 1853a: 147. Type-species: Acrocoelia ruficeps (junior synonym of Formica scutellaris), by subsequent designation of Wheeler, W.M. 1911f: 158.
 * Acrocoelia junior synonym of Crematogaster: Mayr, 1863: 404; Wheeler, W.M. 1911f: 158; Wheeler, W.M. 1922a: 828.
 * Acrocoelia revived from synonymy as subgenus of Crematogaster: Emery, 1922e: 140.
 * Acrocoelia junior synonym of Crematogaster: Buren, 1959: 126.
 * Acrocoelia revived from synonymy as genus: Soulié, 1964: 398.
 * Acrocoelia junior synonym of Crematogaster: Brown, 1973b: 178; Blaimer 2012: 52. [The type-species of Acrocoelia and Crematogaster are synonymous, the generic synonymy is therefore absolute.]
 * ATOPOGYNE [junior synonym of Crematogaster (Crematogaster)]
 * Atopogyne Forel, 1911d: 343 [as subgenus of Crematogaster]. Type-species: Crematogaster (Atopogyne) hellenica, by subsequent designation of Wheeler, W.M. 1911f: 159.
 * [Type-species not Crematogaster depressa, unjustified subsequent designation by Emery, 1912d: 272; this error repeated in Emery, 1922e: 154 and in Soulié, 1965: 78.]
 * Atopogyne raised to genus: Soulié, 1964: 398.
 * Atopogyne junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Atopogyne subgenus of Crematogaster: Bolton, 1995b: 21.
 * Atopogyne junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * COLOBOCREMA [junior synonym of Crematogaster (Crematogaster)]
 * Colobocrema Wheeler, W.M. 1927f: 31 [as subgenus of Crematogaster]. Type-species: Crematogaster (Colobocrema) cylindriceps, by monotypy.
 * Colobocrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Colobocrema subgenus of Crematogaster: Bolton, 1995b: 24.
 * Colobocrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * DECACREMA [junior synonym of Crematogaster (Crematogaster)]
 * Decacrema Forel, 1910a: 18 [as subgenus of Crematogaster]. Type-species: Crematogaster schencki, by subsequent designation of Wheeler, W.M. 1911f: 161.
 * [Decacrema also described as new by Forel, 1910f: 9.]
 * Decacrema raised to genus: Soulié, 1964: 398.
 * Decacrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Decacema subgenus of Crematogaster: Bolton, 1995b: 26.
 * Decacema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 52.
 * [ Decracrema Arnold, 1920a: 547, incorrect subsequent spelling.]
 * NEMATOCREMA [junior synonym of Crematogaster (Crematogaster)]
 * Nematocrema Santschi, 1918d: 182 [as subgenus of Crematogaster]. Type-species: Crematogaster stadelmanni, by original designation.
 * Nematocrema raised to genus: Soulié, 1964: 398.
 * Nematocrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Nematocrema subgenus of Crematogaster: Bolton, 1995b: 38.
 * Nematocrema junior synonym of Oxygyne: Blaimer, 2012: 249.
 * Nematocrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * OXYGYNE [junior synonym of Crematogaster (Crematogaster)]
 * Oxygyne Forel, 1901e: 375 [as subgenus of Crematogaster]. Type-species: Crematogaster (Oxygyne) daisyi, by subsequent designation of Wheeler, W.M. 1911f: 169.
 * Oxygyne raised to genus: Soulié, 1964: 398.
 * Oxygyne junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Oxygyne subgenus of Crematogaster: Bolton, 1995b: 40.
 * Oxygyne senior synonym of Nematocrema: Blaimer, 2012: 249.
 * Oxygyne junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 52.
 * PARACREMA [junior synonym of Crematogaster (Crematogaster)]
 * Paracrema Santschi, 1918d: 182 [as subgenus of Crematogaster]. Type-species: Crematogaster spengeli, by original designation.
 * Paracrema raised to genus: Soulié, 1964: 398.
 * Paracrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Paracrema subgenus of Crematogaster: Bolton, 1995b: 41.
 * Paracrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * PHYSOCREMA [junior synonym of Crematogaster (Crematogaster)]
 * Physocrema Forel, 1912f: 220 [as subgenus of Crematogaster]. Type-species: Crematogaster inflata, by subsequent designation of Wheeler, W.M. 1913a: 82.
 * Physocrema raised to genus: Soulié, 1964: 398.
 * Physocrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Physocrema subgenus of Crematogaster: Bolton, 1995b: 42.
 * Physocrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * SPHAEROCREMA [junior synonym of Crematogaster (Crematogaster)]
 * Sphaerocrema Santschi, 1918d: 182 [as subgenus of Crematogaster]. Type-species: Crematogaster kneri, by original designation.
 * Sphaerocrema raised to genus: Soulié, 1964: 398.
 * Sphaerocrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Sphaerocrema subgenus of Crematogaster: Bolton, 1995b: 46.
 * Sphaerocrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.
 * XIPHOCREMA [junior synonym of Crematogaster (Crematogaster)]
 * Xiphocrema Forel, 1913k: 80 [as subgenus of Crematogaster]. Type-species: Crematogaster tetracantha, by subsequent designation of Emery, 1922e: 138.
 * Xiphocrema raised to genus: Soulié, 1964: 398.
 * Xiphocrema junior synonym of Crematogaster: Hölldobler & Wilson, 1990: 13.
 * Xiphocrema subgenus of Crematogaster: Bolton, 1995b: 50.
 * Xiphocrema junior synonym of Crematogaster (Crematogaster): Blaimer 2012: 53.

Blaimer (2012) provides the following diagnostic definition for this globally distributed subgenus:

1. Petiole shape (in dorsal view) highly variable: moderately flared (Fig. 14a), broadly flared (Fig. 15a), sharply flared (Fig.16a), greatly flared (Fig. 17a), oval (Fig. 18a) or suboval (Fig. 19a), elongate rectangular (Fig. 20), circular (Fig. 21a), or hexagonal (Fig. 22a), but not rectangular or ovo-rectangular (as in Fig. 3 or 4). If petiole elongate-rectangular (Fig. 20) and thus similar to some Orthocrema species (as in Fig. 5), then propodeal spiracle compressed oval.

2. Petiole with dorsoposterior denticles/tubercules and dorsoposterior erect setae (e.g. Figs 14a, 16a) or lacking denticles/tubercules (e.g. Figs 15a, 18a–19a, 20, 22a), while setae may still be present.

3. Postpetiole varying from distinctly (Figs 16b-17b, 19b, 22b) to weakly bilobed (Figs 14b–15b, 18b), with a broad (Fig. 16b) to thin median longitudinal impression (Fig. 23). More rarely postpetiole only with faint posterior impression (Figs 18b, 25) or lacking impression entirely (Fig. 24). Probably ~80% of the species diversity in this clade has a clearly bilobed postpetiole with a distinct median longitudinal impression.

4. Propodeal spiracle often oval or compressed oval (Fig. 26).

5. Occipital carinae variable, but often reduced (Fig. 27).

6. Antennae usually 11-segmented, rarely 10-segmented or (one species) 9-segmented. Antennal scapes of variable length; if antennae distinctly clubbed, then usually club 3-segmented.

7. Head shape highly variable, but often with a straight (Fig. 28) or medially depressed posterior head margin (Fig. 29) and with suboval or subangular corners.