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Odontomachus troglodytes Santschi HNS

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

Type material: Odontomachus haematodes troglodytes Santschi HNS , 1914 : 58 [ 36 ] . Lectotype worker: Kenya , Shimoni cave ( NHMB ), designated by Brown, 1976: 106 [ 2 ] [examined] AntWeb CASENT0101134 . Raised to species Brown, 1976 : 106 [ 1 ] .

Odontomachus haematodus stanleyi Wheeler HNS , 1922 : 102 [ 37 ] . Type worker: DRC ( Zaire ) Stanleyville , 25° 10'E , 0°30'NFeb 1915 , ( AMNH ) [examined] AntWeb CASENT0104653 , CASENT0104654 . Synonymized with troglodytes HNS by Brown, 1976 : 106 [ 1 ] .

Worker measurements: maximum and minimum based on n = 15 from Madagascar: HL 2.23-2.66, HW (across vertex) 1.56- 1.92, HW (across upper eye margin) 1.69-1.98, CI 74-78, EL 0.40-0.47, ML 1.13-1.33, MI 45-54, SL 2.07-2.42, SI 117-127, WL 2.61-3.07. FL 2.28-2.65, PW 1.02-1.19.

The specimens from Madagascar are notably smaller than specimens in CAS collection from South Africa , ceIntral Africa and Sao Tome. Maximum and minimum measurements based on n = 5: HL 2.52-2.94, HW (across vertex) 1.81-2.25, HW (across upper eye margin) 1.94-2.31, CI 74-79, EL 0.41-0.51, ML 1.19- 1.38, MI 47-49, SL 2.24-2.53, SI 110-122, WL 2.88-3.23. FL 2.42-2.91, PW 1.13-1.36.

Queen measurements: maximum and minimum based on n = 5 from Madagascar: HL 2.59-2.74, HW (across vertex) 1.99- 2.19, HW (across upper eye margin) 2.05-2.18, CI 78-79, EL 0.56-0.59, ML 1.39-1.44, MI 52-55, SL 2.36-2.52, SI 112-119, WL 3.18-3.49. FL 2.67-2.76.

Male measurements: maximum and minimum based on n = 5 from Madagascar: HL 1.00-1.04, HW 1.30-1.35, CI 127- 133, EL 0.68-0.70, SL 0.22-0.26, SI 17-19, WL 2.52-2.59. FL 1.80-1.88

Worker Diagnosis: Workers of this species can be easily distinguished from coquereli HNS by their smaller size, distinct extraocular furrows and temporal ridges on vertex and short and blunt mandibular teeth. Brown ( 1976 ) provides additional description and references.

Distribution and biology. O. troglodytes HNS was first reported from Madagascar by Andre [ 38 :290] as O. haematodes (Linnaeus) HNS . African and Malagasy records of haematodes HNS actually refer to troglodytes HNS . In Madagascar, troglodytes HNS is widespread throughout the east in secondary habitats, including coastal scrub, eucalyptus plantations , littoral forest, and rainforest below 800 m elevation. This species is also widespread across sub-Saharan Africa in second growth forests and open habitats. Forel [ 25 :159] recorded Odontomachus HNS (as haematodes HNS ) from Seychelles. These specimens have not been examined but probably refer to O. simillimus HNS and not troglodytes HNS .

Figure 13. Odontomachus HNS spp. ventral aspect of posterior mesosoma viewed from underneath and from rear with coxa and petiole removed to show metasternal process. A, troglodytes HNS CASENT0009961. B, simillimus HNS CASENT0009988. C, coquereli HNS CAS- NET0009962. doi:10.1371/journal.pone.0001787.g013

Figure 14. collection localities of Odontomachus HNS in Madagascar. Map shows major ecoregions: east (light gray): rainforest, ceIntral (dark gray): montane forest; west (white): tropical dry forest; southwest (medium gray): desert spiny bush thicket. doi:10.1371/journal.pone.0001787.g014

Figure 15. NJ tree of K2P for three species of Odontomachus HNS in Madagascar and Africa (all specimens with 500 bp). Deep divergences evident between coquereli HNS , troglodytes HNS , and simillimus HNS are evident. Deep divergences within O. coquereli HNS are apparent. The rightmost column of colors differentiate which biogeographical groupings of Wilme et al [ 29 ] these populations fall. WCE-1 = Binara. WCE-10 = Manongarivo. WCE-2 = Mahavelona, Kalalao, Betampona, Mananara-Nord, Marojejy, Anjanaharibe. WRDW-a2 = Akirindro, Ambanitaza, Anjanaharibe. doi:10.1371/journal.pone.0001787.g015

Figure 16. Anochetus HNS spp. CO1 DNA barcode heterogeneity. A. grandidieri HNS (n = 113), A. madagascarensis HNS (n = 115), A. goodmani HNS (n = 47), A. boltoni HNS (n = 12) and A. pattersoni HNS (n = 3). doi:10.1371/journal.pone.0001787.g016

Because of its preference of secondary habitats, it is possible that troglodytes HNS in Madagascar is a recent colonist from Africa, possibly introduced by humans. This is in coIntrast to coquereli HNS which is most closely related to Melanesian species in the tyrannicus group.

Our collections in Madagascar were focused primarily on less disturbed habitats, thus the distribution map (Fig. 10b) probably does not reflect the full extent of its range. O. troglodytes HNS was most often recorded nesting in rotten logs (30 collection records) followed by sifted litter (15). Males were collected at light, malaise traps, and yellow pan traps.

A lab colony was kept for a number of months and thrived on a diet of crickets, producing numerous larvae, brood, and males. The trap jaw behavior is very similar to that of O. bauri HNS [ 39 , Fisher unpublished]. When disturbed, the specimen use trap jaw propulsion to "jump" away.

CO1. Shallow iIntraspecific and deep interspecific divergences between O. troglodytes HNS in Madagascar and Africa and the other species - what one might expect if it has been recently introduced. Average within species sequence divergence of 0.4% (Figs 15, 17).

Diagnostic barcoding loci. O. troglodytes HNS : G- 1659 , G-465, G- 519, T-535, A-537.

Specimens examined for Odontomachus troglodytes HNS :

Specimens from 105 separate collection events from the following 40 localities were examined. CAMEROON : Sud : Res. de Faune de Campo , 2.16 km 106° ESE Ebodje ; Sud-Ouest : Bimbia Forest , 7.4 km 119° ESE Limbe . CENTRAL AFRICAN REPUBLIC : Prefecture Sangha-Mbaere : Parc National Dzanga-Ndoki , 39.6 km 174° S Lidjombo ; Parc National Dzanga-Ndoki , 38.6 km 173° S Lidjombo ; Parc National Dzanga-Ndoki , 37.9 km 169° S Lidjombo ; Reserve Speciale de Foret Dense de Dzanga-Sangha , 12.7 km 326° NW Bayanga ; Parc National Dzanga-Ndoki , Mabea Bai , 21.4 km 53° NE Bayanga . GABON : Estuaire : Pointe Ngombe , Ekwata , 16 km 240° WSW Libreville ; Libreville ; F.C. Mondah , 21 km 331° NNW Libreville . GABON : Ogooue-Maritime : Aire d'Exploit . Rationnelle de Faune des Monts Doudou , 25.2 km 304° NW Doussala ; Reserve de la Moukalaba-Dougoua , 12.2 km 305 NW Doussala ; Reserve de Faune de la Moukalaba-Dougoua, 12.2 km 305° NW Doussala ; Reserve de Faune de la Moukalaba-Dougoua, 10.8 km 214° SW Doussala ; Woleu-Ntem : 31.3 km 108° ESE Minvoul ; KENYA : [Cote d' Afrique or. angl. Shimoni ; LIBERIA : Sapo Nat. Park . MADAGASCAR : Toamasina : Mahavelona (= Foulpointe ); 5.3 km SSE Ambanizana , Andranobe ; Foret d'AnalavaMandrisy , 5.9 km 195° Antanambe ; Res. Ambodiriana , 4.8 km 306° Manompana , along Manompana river ; Ile Sainte Marie , Foret Ambohidena , 22.8 km 44° Ambodifotatra ; Ile Sainte Marie , Foret Ampanihy , 14.4 km 52° Ambodifotatra ; Ile Sainte Marie , Foret Kalalao , 9.9 km 34° Ambodifotatra ; Parcell K9 Tampolo ; Tampolo ; S.F. Tampolo, 10 km NNE Fenoarivo Atn . ; Parcelle E3 Tampolo ; Parcelle K7 Tampolo ; Bridge at Onibi , NW of Mahavelona ; Mahavelona ( Foulpointe ); 2.1 km 315° Mahavelona ; Toamasina ( Tamatave ); Prison de Tamatave ; Station forest de Tampolo , 10 km N Fenerive ; Res. Betampona , Ambodiriana 45 km NW Toamasina ; 10k N Brickaville ; 11 km SE Ampasimanolotra (= Brickaville ); Fianarantsoa : Riv: Ranomafana Aff. de laroka ; Local : Ranomafana RN2 ; Riv: laroka Aff de Rianila;Local : Manakana ; Riv : Mahatsara Aff de Rianila ; Local : Piste vers Brickaville ; Riv : Rongaronga ; Local : Ambodifaho ; Riv : Rianila ( Ivohitra ) ; Local : Antseranambe ; Riv : Santaravina ; Local : Ampasipotsy-pont routier ; Riv : Sandragniro ; Local : Tanambao-Pont routier ; Riv : Farimbogna ; Local : Village 202 ( Pont routier RN2 ) ; Riv : Ilazana ; Local : Gri-gri ; 8k E Kianjavato Vatovavy Forest ; Ranomafana Nat. Park ; 10k E Ranomafana ; Ranomafana Nat. Park , 10 km E; Mananjary 2 km south; 7.6 km 122° Kianjavato , Foret Classee Vatovavy ; SOUTH AFRICA : Mpumalanga : Songimuelo Nat. Reserve , Kromdraai Camp , Komati River ; Natal : Mtunzini ; Limpopo : Dunstable Farm , 27 km E ofHoedspruit . DEMOCRATIC REPUBLIC OF THE CONGO : Stanleyville ; Epulu .

Complementary analyses to CO1

In some instances we chose to amplify independent nuclear markers to help interpret CO1 divergences involving populations where specimens were morphologically cryptic. Because of their high copy number and relatively conserved primer regions, we selected three ribosomal regions to amplify: 18S , 28S and ITS1. We had high expectations for the utility of these markers to complement the mtDNA barcode analysis based on our own experiences with other taxa [40,41], the utility of these markers in other taxonomic groups where, for instance, ITS1 functions as a barcode [ 42 ], and, for 28S , based on predictions of others for the utility of this region as an alternative barcode region [ 43 ]. Unfortunately, we found that, while the CO1 data from species with exclusively (putatively) ergatoid queens had large phylogeographic signal, when compared to the three rRNA regions we utilized it was markedly simpler to generate, interpret and analyze. The rRNA markers utilized here, particularly 18S and 28S , can be useful for identifying interspecific (species as revised here) hybridization [see 40,41,43].

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