Comprehensive Description

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Anochetus goodmani Fisher HNS ZBK sp. nov. 76BCF3886010

Figures: worker 2e,f; queen 2g,h; map 6a

Type Material: Holotype worker, MADAGASCAR , Foret de Binara , 7.5 km 230° SW Daraina , 13°15'18"S , 049°37'00"E , 375 m, 1-4 Dec 2003 (coll. B. L. Fisher et al.), collection code: BLF09638, pin code: CASENT0498309 ( CAS ). Paratypes : 8 workers with same data as holotype but pins coded, CASENT104548 , CASENT0498310 , CASENT0498311 , CASENT0006944 , CASENT0006945 ( BMNH , MCZ , CAS ) ; CO1 Barcode from paratype collection and coded CASENT0498310-D01 .

Worker measurements: maximum and minimum based on all specimens, n= 15, (holotype): HL 1.77-2.01 (1.92), HW 1.55- 1.81 (1.77), CI 86-92 (92), EL 0.35-0.43 (0.42), ML 1.04-1.15 (1.11), MI 56-66 (58), SL 1.68-1.97 (1.79) SI 101-109 (101), WL 2.52-2.89 (2.66), FL 1.85-2.17 (2.03), PW 0.92-1.06 (1.01).

Queen (ergatoid) measurements: maximum and minimum based on n = 5. HL 1.62-1.79, HW 1.49-1.65, CI 91-93, EL 0.37-0.41, ML 0.92-1.02, MI 55-59, SL 1.56-1.71, SI 99- 106, WL 2.33-2.55, FL 1.77-1.91, PW 0.88-0.99.

Worker Diagnosis: Blade of mandible with five teeth and denticles located at the distal halfofthe blade length. Petiole dorsal margin without spines. In front view, the dorsal petiolar margin flat with lateral margin rounded (Fig. 6b). Pilosity, sculpture as in Figures 2e,f.

The species is most similar to A. boltoni HNS but can be easily distinguished by its petiole node without apical spines.

No winged queens are known. Ergatoid queens were collected at six localities. In four of the collections, three ergatoid queens were collected in the same locality. They are very similar in size and shape to workers (Figs 2g,h), and have no ocelli (Fig. 2g). Males are not known.

Distribution and biology. A. goodmani HNS is endemic to Madagascar and is widespread in northern and western parts of the island. It has been collected in dry forest and rainforest as low as 30 m in altitude and also in montane rainforest at the altitude 960 m on Montagne d'Ambre (Fig. 6a), most frequently under stones (12 collections) and sifted litter (7), but also at light (1), beating low vegetation (3), rot pocket (1), in rotten log (6), ground foragers (1), ground nest (9), Malaise trap (1), on low vegetation (1), and pitfall traps (4).

Figure 6. collection localities of Anochetus HNS specimens 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.g006

Figure 7. Anochetus pattersoni HNS . A-D Worker holotype CASENT0102280 full face, lateral view, upper part of petiole from rear view, dorsal view. E-F, queen paratype CASENT0103343 full face and lateral view. G-H, male CASENT0172617 full face and lateral view. doi:10.1371/journal.pone.0001787.g007

CO1. Average iIntraspecific sequence divergence of 6.37%. There is strong geographic coherence in the divergence patterns (Figs 9, 15, Table 2) with deep divergences occurring between separate regions isolated by habitat and mountains.

Diagnostic barcoding loci. A. goodmani HNS : Y-231 ( madagascarensis HNS and grandidieri HNS A; boltoni HNS and pattersoni HNS T), W-233 (all others A), RWR-368-370 (others are all ATG), Y-541 (others are all T), R-543 (others are all A), W-546 (others are all T), W-585 (others are all T), M-634 (others are all C). RWCW-42-45 & WTTAG-66-70 (this distinguishes goodmani HNS from all (including boltoni HNS ) except some madagascarensis HNS ), & GT-83-84 ( madagascarensis HNS is TA).

Discussion. Anochetus goodmani HNS is characterized by extreme divergence within the barcode region. To date, sequencing complementary nuclear markers has provided some degree of support for the deepest CO1 divergences (between the north and south-west of Madagascar) as being separate species. Importantly however, ITS1 sequences as divergent have been produced from the same individual (Appendix S1 and Table 3). Although CO1 supports more than one operational unit within A. goodmani HNS the hypothesis of cryptic species in relatively isolated environments requires further evidence with less ambiguity.

Additional material examined for Anochetus goodmani HNS : In addition to the type material, specimens from 56additional collecting events from the following 18 localities were examined in this study. MADAGASCAR : Province Antsiranana : Montagne des Francais , 7.2 km 142° SE Antsiranana ; Parc National Montagne d'Ambre ; Reserve Speciale de l'Ankarana , 13.6 km 192° SSW Anivorano Nord ; Reserve Speciale de l'Ankarana , 22.9 km 224° SW Anivorano Nord ; Foret d'Ampondrabe, 26.3 km 10° NNE Daraina ; Foret d' Andavakoera , 21.4 km 75° ENE Ambilobe ; 4.6 km 356° NBetsiaka ; Foret d' Antsahabe , 11.4 km 275° WDaraina ; Foret de Binara , 7.5 km 230° SW Daraina ; Ampasindava, Foret d'Ambilanivy , 3.9 km 181° SAmbaliha ; Foret d'Anabohazo , 21.6 km 247° WSW Maromandia ; Reserve Speciale de Bemarivo , 23.8 km 223° SW Besalampy ; Parc National Tsingy de Bemaraha , 10.6 km ESE 123° Antsalova ; Parc National Tsingy de Bemaraha , 2.5 km 62 ° ENE Bekopaka , Ankidrodroa River ; Parc National Tsingy de Bemaraha , 3.4 km 93° EBekopaka , Tombeau Vazimba . Province Toliara : Parc National de Kirindy Mite , 16.3 km 127° SE Belo sur Mer .

Figure 8. Anochetus HNS males, terminalia, lateral view. A, boltoni HNS CASENT0063847. B, grandidieri HNS CASENT0080660. B, madagascarensis HNS CASENT0063421. D, pattersoni HNS CASENT0172617. doi:10.1371/journal.pone.0001787.g008


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