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Species: Odontomachus troglodytes   Santschi, 1914 

Classification:
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Taxonomic History (provided by Barry Bolton, 2016)

Odontomachus haematodes var. troglodytes Santschi, 1914b: 58 (w.) KENYA. Afrotropic. "Kenya, Shimoni cave." Naturhistorisches Museum, Basel, Switzerland (NHMB). CASENT0101134. http://www.antweb.org/specimen/CASENT0101134 AntCat AntWiki HOL

Taxonomic history

André, 1887 PDF: 290 (m.); Arnold, 1915: 109 (q.); Menozzi, in Eidmann, 1944: 437 (l.).
Raised to species and senior synonym of Odontomachus stanleyi: Brown, 1976a: 106 (Lectotype designation).
See also: Brown, 1976a}: 167; Fisher & Smith, 2008 PDF: 16.

Distribution:


Afrotropical Region: Benin, Botswana, Cameroon, Central African Republic, Congo, Democratic Republic of Congo, Equatorial Guinea, Eritrea, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Mozambique, Sao Tome and Principe, Senegal, South Africa, Tanzania, Uganda, Zambia, Zimbabwe
Malagasy Region: Madagascar

Distribution Notes:

Reference for Kenya if not type: Kakamega Forest

Notes:

Caste of types: (w)

Taxonomic Treatment (provided by Plazi)

Treatment Citation: Fisher, B. L. & Smith, M. A., 2008, A revision of Malagasy species of Anochetus Mayr and Odontomachus Latreille (Hymenoptera: Formicidae)., PLoS ONE 3, pp. 1-23

Odontomachus troglodytes SantschiHNS

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

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

Odontomachus haematodus stanleyi WheelerHNS, 1922: 102 [37]. Type worker: DRC ( Zaire) Stanleyville GoogleMaps, 25° 10'E, 0°30'N Feb 1915, (AMNH) [examined] AntWeb CASENT0104653 GoogleMapsCASC, CASENT0104654. GoogleMapsCASC Synonymized with troglodytesHNS 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 coquereliHNS 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. troglodytesHNS was first reported from Madagascar by Andre [38:290] as O. haematodes (Linnaeus)HNS. African and Malagasy records of haematodesHNS actually refer to troglodytesHNS. In Madagascar, troglodytesHNS 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 OdontomachusHNS (as haematodesHNS) from Seychelles. These specimens have not been examined but probably refer to O. simillimusHNS and not troglodytesHNS.

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

Figure 14. collection localities of OdontomachusHNS 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 OdontomachusHNS in Madagascar and Africa (all specimens with 500 bp). Deep divergences evident between coquereliHNS, troglodytesHNS, and simillimusHNS are evident. Deep divergences within O. coquereliHNS 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. AnochetusHNS spp. CO1 DNA barcode heterogeneity. A. grandidieriHNS (n = 113), A. madagascarensisHNS (n = 115), A. goodmaniHNS (n = 47), A. boltoniHNS (n = 12) and A. pattersoniHNS (n = 3). doi:10.1371/journal.pone.0001787.g016

Because of its preference of secondary habitats, it is possible that troglodytesHNS in Madagascar is a recent colonist from Africa, possibly introduced by humans. This is in coIntrast to coquereliHNS 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. troglodytesHNS 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. bauriHNS [39, Fisher unpublished]. When disturbed, the specimen use trap jaw propulsion to "jump" away.

CO1. Shallow iIntraspecific and deep interspecific divergences between O. troglodytesHNS 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. troglodytesHNS: G- 1659, G-465, G- 519, T-535, A-537.

Specimens examined for Odontomachus troglodytesHNS:

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 GoogleMaps); 5.3 km SSE Ambanizana GoogleMaps, Andranobe GoogleMaps; GoogleMapsCASC Foret d'Analava Mandrisy , 5.9 km 195° Antanambe ; Res. Ambodiriana, 4.8 km 306° Manompana GoogleMaps, along Manompana river GoogleMapsGoogleMapsCASCCASC1GB1; Ile Sainte Marie, Foret Ambohidena GoogleMaps, 22.8 km 44° Ambodifotatra GoogleMapsGoogleMapsCASCCASC1GB1; Ile Sainte Marie, Foret Ampanihy GoogleMaps, 14.4 km 52° Ambodifotatra GoogleMaps; GoogleMapsCASCCASC1GB1 Ile Sainte Marie, Foret Kalalao GoogleMaps, 9.9 km 34° Ambodifotatra GoogleMapsGoogleMapsCASCCASC1GB1; Parcell K9 Tampolo ; Tampolo GoogleMapsGoogleMapsCASC; S.F. Tampolo, 10 km NNE Fenoarivo Atn GoogleMaps. GoogleMapsCASCCASC1GB1; Parcelle E3 Tampolo GoogleMapsGoogleMapsCASC; Parcelle K7 Tampolo GoogleMaps; GoogleMapsCASC Bridge at Onibi, NW of Mahavelona GoogleMapsGoogleMapsCASC; Mahavelona ( Foulpointe GoogleMaps); 2.1 km 315° Mahavelona GoogleMaps; GoogleMapsCASCCASC1CASC2GB1GB2 Toamasina ( Tamatave GoogleMaps); Prison de Tamatave GoogleMaps; GoogleMapsCASC Station forest de Tampolo, 10 km N Fenerive GoogleMapsGoogleMaps; Res. Betampona, Ambodiriana GoogleMaps 45 km NW Toamasina GoogleMapsGoogleMaps; 10k N Brickaville GoogleMapsGoogleMapsCASC; 11 km SE Ampasimanolotra GoogleMaps (= Brickaville GoogleMaps); GoogleMapsCASC Fianarantsoa: Riv: Ranomafana GoogleMaps Aff. de laroka GoogleMaps; GoogleMapsCASC Local: Ranomafana GoogleMaps RN2 GoogleMapsCASC; Riv: laroka Aff de Rianila; Local: Manakana ; Riv: Mahatsara Aff de Rianila ; Local: Piste vers Brickaville GoogleMapsGoogleMapsCASC; 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 GoogleMapsGoogleMapsCASC; Ranomafana GoogleMaps Nat. Park GoogleMaps; 10k E Ranomafana GoogleMapsGoogleMapsCASC; Ranomafana Nat. Park, 10 km E; Mananjary GoogleMaps 2 km south; 7.6 km 122° Kianjavato GoogleMaps, Foret Classee Vatovavy GoogleMaps; GoogleMapsCASC 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].

Specimen Habitat Summary

Found most commonly in these habitats: 33 times found in rainforest, 22 times found in littoral rainforest, 15 times found in urban/garden, 13 times found in urban garden, 15 times found in littoral forest, 6 times found in coastal scrub, 4 times found in Lydenburgia - Euclea Open Woodland (in Sekhukhune Mountain Bushveld), 2 times found in eucalyptus plantation, 1 times found in low second growth, 2 times found in Lydenburgia - Vitex Open Woodland on hillslope (in Sekhukhune Mountain Bushveld), ...

Found most commonly in these microhabitats: 36 times ex rotten log, 14 times ground forager(s), 14 times sifted litter (leaf mold, rotten wood), 7 times under rotten log, 6 times ex root mat, ground layer, 5 times ground nest, 4 times on low vegetation, 1 times on ground, 2 times ex rot pocket above ground, 2 times ex dead twig above ground, 1 times nest in underground stump near seashore, ...

Collected most commonly using these methods: 9 times taken at light at night, 9 times GC, 10 times MW 50 sample transect, 5m, 9 times hand collected, 6 times pitfall trap, 4 times pitfall trap, PF bucket, 5 times Malaise trap, 4 traps, 5 times EC31 yellow pan trap, 1 times search, 3 times MW, 10 sifted litter, 2 times ex yellow pan trap, ...

Elevations: collected from 5 - 1677 meters, 230 meters average

Type specimens: Lectotype of Odontomachys troglodytes: casent0101134; paratype of Odontomachus haematodus stanleyi: rmcaent000017743; syntype of Odontomachus haematodus stanleyi: casent0104653, casent0104654, casent0915246



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