Odontomachus coquereli Roger 1861

Colonies (N=29) consisted of 19±11 workers and one ergatoid (permanently wingless) queen (Molet et al. 2007). All other species in the genus have winged queens.

Odontomachus coquereli Roger, 1861a PDF: 30 (w.) MADAGASCAR. Malagasy. Primary type information: Madagascar, coll. Coquerel; CASENT0104549; ZMHB AntCat AntWiki HOL

Taxonomic history

Fisher & Smith, 2008 10.1371/journal.pone.0001787 PDF: 14 (q.m.).Combination in Stenomyrmex: Mayr, 1865 PDF: 63 (footnote).Combination in Champsomyrmex: Emery, 1892f PDF: 558.Status as species: Roger, 1863b PDF: 21; Mayr, 1863a PDF: 436; Forel, 1891c PDF: 105 (redescription); Dalla Torre, 1893 PDF: 49; Wasmann, 1897a PDF: 250; Emery, 1911e PDF: 111; Wheeler, 1922: 1013; Brown, 1976c PDF: 103, 143; Bolton, 1995b: 295; Fisher & Smith, 2008 10.1371/journal.pone.0001787 PDF: 13 (redescription).Senior synonym of Odontomachus coquereli minor: Brown, 1976c PDF: 103; Bolton, 1995b: 295; Fisher & Smith, 2008 10.1371/journal.pone.0001787 PDF: 14.

Figures: worker 1b, 10a,b, 13c; queen 10c,d; male 11a,b,e; map 14a

Worker measurements: maximum and minimum based on n = 45 from Madagascar: HL 2.69-3.27, HW (across vertex) 1.26- 1.77, HW (across upper eye margin) 1.54-2.02, CI 57-67, EL 0.46-0.55, ML 1.76-2.16, MI 61-68, SL 3.04-3.96, SI 164-207, WL 4.18-5.11. FL 3.32-4.68, PW 1.11-1.53.

Queen measurements: maximum and minimum based on n = 5 from Madagascar: HL 2.81-2.94, HW (across vertex) 1.39- 1.55, HW (across upper eye margin) 1.83-1.98, CI 62-71, EL 0.45-0.55, ML 1.66-1.81, MI 59-62, SL 3.07-3.29, SI 155-179, WL 4.35-4.56, FL 3.60-3.84, PW 1.28-1.43. Preapical teeth count 7-10.

Male measurements: maximum and minimum based on n = 5 from Madagascar: HL 1.11-1.22, HW 1.41-1.57, CI 128- 134, EL 0.78-0.90, SL 0.30-0.38, SI 21-23, WL 3.38-3.85, FL 2.90-3.16.

Worker Diagnosis: Workers of this species can be easily distinguished from troglodytes by their larger size, mandible with long, acute apical and preapical teeth and lack of extraocular furrows and temporal ridges on vertex. Brown [2] provides a description and additional references.

Distribution and biology. O. coquereli is endemic to Madagascar and is restricted to eastern and northern montane rainforest, lowland rainforest, and littoral forest from 10 to 1325 m (Fig. 10a). It is most abundant at mid-elevations in the northeast such as in Marojejy National Park. Nests of O. coquereli are most commonly found in rotten logs and consist of small colonies. Queens of coquereli are wingless and very similar to workers; colonies reproduce by fission [28]. Males are collected in Malaise traps and yellow pan traps. Workers forage on the ground day and night. A few times BLF has seen solitary foragers high up on trunks and branches of large trees. It is not clear if they are foraging for plant or insect liquids up in the canopy.

There is notable geographic variation in shape of petiole, sculpture and number of preapical teeth. Preapical teeth and denticles range from 7-12. Occasionally, adjacent teeth may be fused at base to form a single bidententate tooth. However, there is no consistent concordant pattern to this variation. Molecular data are also extremely variable - suggesting that these isolated populations have long been separated. Rather than describing these populations as distinct species, we leave them here as a single species - a hypothesis that can be tested in the future with subsequent experiments in both the field and lab.

CO1: The barcode region is extremely variable (Fig. 16) - there is evident isolation by distance which is largely concordant with the biogeographic regions proposed by Wilme et al. [29].

Diagnostic barcoding loci. O. coquereli : T-96, C-196, T-211, T-280, A-283.

Discussion: Odontomachus coquereli from Madagascar, the only species in the genus where winged queens have never been found.

Molet et al. [28] investigated the Marojejy population of O. coquereli , and based on demography, morphometry, allometry and ovarian dissections demonstrated that the winged queen caste has been replaced by a wingless reproductive caste and that the strategy of colonial reproduction is fission. A single wingless reproductive (ergatoid) was found in each colony. The smallest colonies consisted of at least 5 workers and the largest colonies never exceeded 40 workers, indicating a threshold size at which a colony divides in two daughter colonies. In coIntrast, O. troglodytes reproduces by non-claustral independent foundation and colonies can reach 1300 workers [30]. As in A. goodmani and A. boltoni , the other species without winged queens - there are deep CO1 divergences between different collection localities.

Specimens examined for Odontomachus coquereli : Specimens from 134 separate collection events from the following 57 localities were examined.MADAGASCAR : Province Antsiranana : Foret de Binara , 9.4 km 235° SW Daraina ;R.S. Manongarivo , 12.8 km 228° SW Antanambao ;R.S. Manongarivo , 14.5 km 220° SW Antanambao ;RNI Marojejy, 8 km NW Manantenina ;Parc National de Marojejy , Manantenina River ,27.6 km 35° NE Andapa ;Parc National de Marojejy , Manantenina River , 28.0 km 38° NE Andapa ;Parc National de Marojejy , AIntranohofa , 26.6 km 31° NNE Andapa ;Foret Ambanitaza , 26.1 km 347° Antalaha ;Res. Anjanaharibe-Sud , 6.5 km SSW Befingotra , ;Res. D' Anjanaharibe-Sud , 17 km W Andapa ;Province Toamasina : 6.9 km NE Ambanizana , Ambohitsitondroina ;Montagne d'Anjanaharibe , 19.5 km 27° NNE Ambinanitelo ;Montagne d'Anjanaharibe , 18.0 km 21 ° NNE Ambinanitelo ;Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo ;Parc National Masoala , Ambanizana ,;5.3 km SSE Ambanizana , Andranobe ;1 km W Andampibe , Cap Masoala ;Parc National Mananara-Nord , 7.1 km 261 ° Antanambe ;Res. Ambodiriana , 4.8 km 306° Manompana , along Manompana river ;Ile Sainte Marie , Foret Kalalao, 9.9 km 34° Ambodifotatra ;Parcelle E3 Tampolo ;Mahavelona ( Foulpointe ) ;Mahavelona ( Foulpointe ), Forest Andalava ;Reserve Betampona , Camp Vohitsivalana , 37.1 km 338° Toamasina ;Reserve Betampona , Camp Rendrirendry 34.1 km 332° Toamasina ;F.C. Andriantantely ;6 km ESE Andasibe (= Perinet) ;Province Fianarantsoa : Nat. Pk.Ranomafana, Miaranony Forest ;Ranomafana Nat. Park , Valoloaka forest ;Foret d'Ambalagoavy Nord , Ikongo , Ambatombe ;45 km S. Ambalavao ;Res. Andringitra , 43 km S Ambalavao .