Etymology.

Taxon classification

Animalia

Hymenoptera

Formicidae

Csősz

Sándor

Fisher

Brian L.

Diagnostic survey of Malagasy

Nesomyrmex

species-groups and revision of

hafahafa

group species via morphology based cluster delimitation protocol

Zookeys810201520155261959

10.3897/zookeys.526.6037

Nesomyrmex

spinosus

http://zoobank.org/D3643DB1-75EB-415A-9220-9F255A5FCB21

Csősz & Fisher

sp. n.

Figs 28–30, Table 4

Etymology.

Name “spinosus” refers to the short, strong antero-dorsal spines on the petiolar node.

Type material investigated.

Holotype worker. CASENT0443515, BLF05489; MADGAGASCAR: Prov. Toliara, Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325° NW Amboasary, 24°56'S, 46°13'E [-24.92972 N, 46.20967 E], 65 m, 7.iii.2002 Fisher et al. (CAS CASENT0443515);

Paratypes. 24 workers and three males with the same label data with the holotype under CASENT codes: CASENT0443515, BLF05489, (2w, CAS); CASENT0443516, BLF05489, (3w, CAS); CASENT0443517, BLF05489, (3w, CAS); CASENT0443518, BLF05489, (1w, CAS); CASENT0443519, BLF05489, (1w, CAS); CASENT0443520, BLF05489, (1w, CAS); CASENT0443521, BLF05489, (1w, CAS); CASENT0443522, BLF05489, (1w, CAS); CASENT0443523, BLF05489, (1w, CAS); CASENT0443524, BLF05489, (1w, CAS); CASENT0443525, BLF05489, (1w, CAS); CASENT0443526, BLF05489, (1w, CAS); CASENT0443527, BLF05489, (1w, CAS); CASENT0443530, BLF05489, (1w, CAS); CASENT0443531, BLF05489, (1w, CAS); CASENT0443532, BLF05489, (1w, CAS CASENT0443532); CASENT0443533, BLF05489, (1w, CAS); CASENT0443534, BLF05489, (1w, CAS); CASENT0443535, BLF05489, (1w, CAS); CASENT0443536, BLF05489, (1m, CAS); CASENT0443537, BLF05489, (2m, CAS);

The list of 44 non-type individuals belonging to 26 nest samples of other material investigated is given in Table 1.

Diagnosis.

In key.

Description of workers.

Body color: brown. Body color pattern: body concolorous, only clava darker. Absolute cephalic size: 1021 [935, 1121] µm (n=46). Cephalic length vs. maximum width of head capsule (CL/CWb): 1.056 [0.980, 1.113]. Postocular distance vs. cephalic length (PoOc/CL): 0.374 [0.342, 0.393]. Postocular sides of cranium contour frontal view orientation: converging posteriorly. Postocular sides of cranium contour frontal view shape: broadly convex. Vertex contour line in frontal view shape: slightly concave. Vertex sculpture: main sculpture rugose, ground sculpture areolate. Gena contour line in frontal view shape: feebly convex. Genae contour from anterior view orientation: converging. Gena sculpture: rugo-reticulate with areolate ground sculpture. Concentric carinae laterally surrounding antennal foramen count: present. Eye length vs. absolute cephalic size (EL/CS): 0.239 [0.220, 0.265]. Frontal carina distance vs. absolute cephalic size (FRS/CS): 0.315 [0.291, 0.335]. Longitudinal carinae on median region of frons count: present. Longitudinal carinae on medial region of frons shape: forked. Smooth median region on frons count: absent. Antennomere count: 12. Scape length vs. absolute cephalic size (SL/CS): 0.880 [0.844, 0.919]. Facial area of the scape absolute setal angle: setae absent, pubescence only. Median clypeal notch count: present. Median clypeal notch depth vs. absolute cephalic size (Cdep/CS): 0.021 [0.015, 0.027]. Ground sculpture of submedian area of clypeus: smooth. Median carina of clypeus count: present. Lateral carinae of clypeus count: present. Median anatomical line of propodeal spine angle value to Weber length in lateral view: 65°. Spine length vs. absolute cephalic size (SPST/CS): 0.300 [0.258, 0.330]. Minimum spine distance vs. absolute cephalic size (SPBA/CS): 0.212 [0.184, 0.235]. Apical spine distance vs. absolute cephalic size (SPTI/CS): 0.307 [0.221, 0.361]. Propodeal spine shape: straight; slightly bent. Apical distance of pronotal spines vs. absolute cephalic size (PSTI/CS): 0.677 [0.624, 0.723]. Metanotal depression count: absent. Dorsal region of mesosoma sculpture: rugose with areolate ground sculpture. Lateral region of pronotum sculpture: areolate ground sculpture, superimposed by dispersed rugae. Mesopleuron sculpture: areolate ground sculpture, superimposed by dispersed rugae. Metapleuron sculpture: areolate ground sculpture, superimposed by dispersed rugae. Petiole width vs. absolute cephalic size (PEW/CS): 0.237 [0.206, 0.259]. Anterodorsal spines on petiolar node angle of deviation from each other: 60°. Apical distance of anterodorsal spines on petiolar node vs. absolute cephalic size (NSTI/CS): 0.216 [0.194, 0.276]. Frontal profile of petiolar node contour line in lateral view shape: straight. Dorso-caudal petiolar profile contour line in lateral view shape: convex. Dorsal region of petiole sculpture: ground sculpture areolate, main sculpture absent; ground sculpture areolate, main sculpture dispersed rugose. Postpetiole width vs. absolute cephalic size (PPW/CS): 0.491 [0.435, 0.529]. Dorsal region of postpetiole sculpture: ground sculpture areolate, main sculpture absent; ground sculpture areolate, main sculpture dispersed rugose.

Figures 28–30.

Nesomyrmex spinosus sp. n. paratype worker (CASENT0443532). Lateral view of the body (28), head of the holotype worker in full-face view (29), dorsal view of the body (30). Scale 0.5 mm.

Distribution.

This species is known to occur in small, highly isolated forests (Réserve Privé Berenty, Forêt d’Anjapolo and Parc National d’Andohahela, Forêt d’Ambohibory) in the southern part of Madagascar (Fig. 13).

Table 4.

Morphometric data of species calculated on individuals. Mean of indices, ±SD are provided in the upper row, minimum and maximum values are given in parentheses in the lower row.

Species:	Nesomyrmex capricornis sp. n.	Nesomyrmex hafahafa sp. n.	Nesomyrmex medusus sp. n.	Nesomyrmex spinosus sp. n.
nr. of individulals:	(n = 27)	(n = 48)	(n = 56)	(n = 46)
CS	1024±38	1062±41	1069±52	1021±43
	[919, 1115]	[974, 1142]	[958, 1189]	[935, 1121]
CL/CWb	1.079±0.020	1.038±0.020	1.046±0.025	1.056±0.024
	[1.037, 1.111]	[0.993, 1.075]	[0.990, 1.097]	[0.980, 1.113]
PoOC/CL	0.390±0.006	0.388±0.010	0.391±0.008	0.374±0.011
	[0.381, 0.403]	[0.361, 0.406]	[0.371, 0.413]	[0.342, 0.393]
FRS/CS	0.315±0.007	0.316±0.008	0.313±0.008	0.315±0.009
	[0.297, 0.326]	[0.289, 0.333]	[0.295, 0.331]	[0.291, 0.335]
SL/CS	0.927±0.012	0.895±0.017	0.907±0.028	0.880±0.016
	[0.907, 0.948]	[0.861, 0.927]	[0.849, 0.997]	[0.844, 0.919]
EL/CS	0.241±0.011	0.230±0.007	0.232±0.007	0.239±0.008
	[0.225, 0.267]	[0.212, 0.248]	[0.219, 0.249]	[0.220, 0.265]
MW/CS	0.652±0.012	0.657±0.019	0.682±0.018	0.650±0.014
	[0.632, 0.685]	[0.631, 0.712]	[0.633, 0.740]	[0.618, 0.679]
PEW/CS	0.265±0.017	0.307±0.021	0.268±0.011	0.237±0.009
	[0.238, 0.312]	[0.275, 0.357]	[0.246, 0.295]	[0.206, 0.259]
PPW/CS	0.558±0.025	0.538±0.022	0.543±0.021	0.491±0.022
	[0.516, 0.613]	[0.494, 0.576]	[0.496, 0.585]	[0.435, 0.529]
SPBA/CS	0.260±0.014	0.287±0.014	0.266±0.018	0.212±0.010
	[0.238, 0.292]	[0.257, 0.311]	[0.234, 0.308]	[0.184, 0.235]
SPTI/CS	0.455±0.039	0.543±0.032	0.443±0.034	0.307±0.027
	[0.386, 0.569]	[0.463, 0.607]	[0.354, 0.504]	[0.221, 0.361]
ML/CS	1.290±0.026	1.266±0.029	1.319±0.031	1.270±0.023
	[1.234, 1.335]	[1.201, 1.323]	[1.181, 1.376]	[1.218, 1.313]
PEL/CS	0.506±0.015	0.420±0.014	0.441±0.018	0.435±0.010
	[0.468, 0.526]	[0.399, 0.453]	[0.392, 0.500]	[0.397, 0.459]
NOL/CS	0.303±0.017	0.278±0.015	0.290±0.012	0.299±0.012
	[0.258, 0.338]	[0.229, 0.307]	[0.243, 0.319]	[0.265, 0.321]
PPL/CS	0.216±0.007	0.202±0.010	0.211±0.009	0.206±0.011
	[0.204, 0.228]	[0.181, 0.223]	[0.190, 0.233]	[0.164, 0.231]
SPST/CS	0.397±0.017	0.398±0.019	0.385±0.019	0.300±0.018
	[0.367, 0.432]	[0.355, 0.427]	[0.333, 0.437]	[0.258, 0.330]
MPST/CS	0.411±0.011	0.409±0.013	0.400±0.010	0.404±0.012
	[0.386, 0.432]	[0.383, 0.442]	[0.379, 0.426]	[0.370, 0.433]
PSTI/CS	0.658±0.017	0.724±0.028	0.757±0.020	0.677±0.021
	[0.617, 0.690]	[0.631, 0.776]	[0.711, 0.813]	[0.624, 0.723]
NSTI/CS	0.265±0.035	0.514±0.052	0.354±0.039	0.216±0.018
	[0.203, 0.364]	[0.473, 0.563]	[0.278, 0.464]	[0.194, 0.276]
Cdep/CS	0.023±0.003	0.022±0.003	0.022±0.003	0.021±0.005
	[0.018, 0.030]	[0.015, 0.029]	[0.017, 0.029]	[0.015, 0.027]

Table 1.

List of morphometrically investigated samples. Unique CASENT number for pinned samples, locality, geographic coordinates (E, N) in decimal format altitude (ALT) in meters a.s.l., collector’s name, date and number of specimens investigated bearing the given CASENT number are provided. Red row: holotype, yellow row: paratype(s). All samples collected in Toliara administrative region, Madagascar, and deposited at the California Academy of Sciences (CAS).

Species name	CASENT number	Locality	N	E	ALT	Collector	Date	Number of specimens
capricornis sp. n. HT	CASENT0452741	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	1w
capricornis sp. n. PT	CASENT0452715	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	1w
capricornis sp. n. PT	CASENT0452716	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	1w
capricornis sp. n. PT	CASENT0452738	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	2w
capricornis sp. n. PT	CASENT0452739	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	2w
capricornis sp. n.	CASENT0443010	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	1w
capricornis sp. n.	CASENT0456949	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	2w
capricornis sp. n.	CASENT0456950	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	2w
capricornis sp. n.	CASENT0452881	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	1w
capricornis sp. n.	CASENT0459109	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0459110	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0456620	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0456621	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0452872	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	2w
capricornis sp. n.	CASENT0452175	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	2w
capricornis sp. n.	CASENT0452871	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	2w
capricornis sp. n.	CASENT0020707	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0079196	Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63°ENE Amboasary, 7.6 km 99°E Hazofotsy	-24,817	46,61	150 m	Fisher-Griswold Arthropod Team	1/12/2002	1w
capricornis sp. n.	CASENT0452754	Forêt de Mahavelo, Isantoria River	-24,758	46,157	110 m	Fisher-Griswold Arthropod Team	1/28/2002	3w

hafahafa sp. n. HT	CASENT0460666	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	1w
hafahafa sp. n. PT	CASENT0746771	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	1w
hafahafa sp. n. PT	CASENT0460667	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0430386	Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer	-20,795	44,147	80 m	Fisher-Griswold Arthropod Team	12/6/2001	2w
hafahafa sp. n.	CASENT0430386	Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer	-20,795	44,147	80 m	Fisher-Griswold Arthropod Team	12/6/2001	2w
hafahafa sp. n.	CASENT0430494	Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer	-20,795	44,147	80 m	Fisher-Griswold Arthropod Team	12/6/2001	2w
hafahafa sp. n.	CASENT0430390	Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer	-20,795	44,147	80 m	Fisher-Griswold Arthropod Team	12/6/2001	2w
hafahafa sp. n.	CASENT0451365	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0460712	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0457087	Forêt de Beroboka, 5.9 km 131°SE Ankidranoka	-22,233	43,366	80 m	Fisher-Griswold Arthropod Team	3/12/2002	2w
hafahafa sp. n.	CASENT0439492	Forêt de Beroboka, 5.9 km 131°SE Ankidranoka	-22,233	43,366	80 m	Fisher-Griswold Arthropod Team	3/12/2002	2w
hafahafa sp. n.	CASENT0460679	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0457090	Forêt de Beroboka, 5.9 km 131°SE Ankidranoka	-22,233	43,366	80 m	Fisher-Griswold Arthropod Team	3/12/2002	2w
hafahafa sp. n.	CASENT0426075	3 km 50°NE Ifaty	-23,15	43,617	60 m	D.O.Burge	10/23/2001	2w
hafahafa sp. n.	CASENT0426077	3 km 50°NE Ifaty	-23,15	43,617	60 m	D.O.Burge	10/23/2001	2w
hafahafa sp. n.	CASENT0059254	Ranobe	-23,045	43,615	20 m	Frontier Wilderness Project	1/26/2004	1w
hafahafa sp. n.	CASENT0446254	Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer	-20,795	44,147	80 m	Fisher-Griswold Arthropod Team	12/6/2001	1w
hafahafa sp. n.	CASENT0066346	Mikea Forest, spiny forest, Tulear Province	-22,913	43,482	37 m	R. Harin’Hala	11/27/2001	1w
hafahafa sp. n.	CASENT0427038	Forêt de Beroboka, 5.9 km 131°SE Ankidranoka	-22,233	43,366	80 m	Fisher-Griswold Arthropod Team	3/12/2002	1w
hafahafa sp. n.	CASENT0447426	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0447445	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0447465	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	1w
hafahafa sp. n.	CASENT0127637	48 km ENE Morondava, Kirindy	-20,067	44,65	30 m	B.L.Fisher	4/18/1995	2w
hafahafa sp. n.	CASENT0426078	3 km 50°NE Ifaty	-23,15	43,617	60 m	D.O.Burge	10/23/2001	2w
hafahafa sp. n.	CASENT0430746	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0459595	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0004062	Forêt de Tsinjoriaky, 6.2 km 84°E Tsifota	-22,802	43,421	70 m	Fisher-Griswold Arthropod Team	3/6/2002	2w
hafahafa sp. n.	CASENT0457427	Forêt de Beroboka, 5.9 km 131°SE Ankidranoka	-22,233	43,366	80 m	Fisher-Griswold Arthropod Team	3/12/2002	1w

medusus sp. n. HT	CASENT0455428	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	1w
medusus sp. n. PT	CASENT0746770	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	1w
medusus sp. n.	CASENT0448719	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	2/21/2002	2w
medusus sp. n.	CASENT0449033	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	2/21/2002	2w
medusus sp. n.	CASENT0449105	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	2/21/2002	2w
medusus sp. n.	CASENT0448791	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	2/21/2002	2w
medusus sp. n.	CASENT0448943	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	37308	2w
medusus sp. n.	CASENT0448945	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	37308	2w
medusus sp. n.	CASENT0451410	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	37308	2w
medusus sp. n.	CASENT0455001	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	37333	2w
medusus sp. n.	CASENT0448723	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	37308	1w
medusus sp. n.	CASENT0424306	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	37337	1w
medusus sp. n.	CASENT0445085	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	37337	1w
medusus sp. n.	CASENT0444985	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	37337	3w
medusus sp. n.	CASENT0445705	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0445292	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0445591	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0444997	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0427243	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0455177	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	2w
medusus sp. n.	CASENT0445705	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	37337	2w
medusus sp. n.	CASENT0445590	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	2w
medusus sp. n.	CASENT0445291	Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81°E Efoetse, 23.0 km 131°SE Beheloka	-23,992	43,881	90 m	Fisher-Griswold Arthropod Team	3/22/2002	4w
medusus sp. n.	CASENT0004002	Mahafaly Plateau, 6.2 km 74°ENE Itampolo	-24,654	43,997	80 m	Fisher-Griswold Arthropod Team	2/21/2002	2w
medusus sp. n.	CASENT0477179	Parc National de Tsimanampetsotsa, 6.7 km 130°SE Efoetse, 23.0 km 175°S Beheloka	-24,101	43,76	25 m	Fisher-Griswold Arthropod Team	3/18/2002	1w
medusus sp. n.	CASENT0477180	Parc National de Tsimanampetsotsa, 6.7 km 130°SE Efoetse, 23.0 km 175°S Beheloka	-24,101	43,76	25 m	Fisher-Griswold Arthropod Team	3/18/2002	1w
medusus sp. n.	CASENT0455436	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	2w
medusus sp. n.	CASENT0454945	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	2w
medusus sp. n.	CASENT0454890	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	2w
medusus sp. n.	CASENT0455002	Parc National de Tsimanampetsotsa, Mitoho Cave, 6.4 km 77°ENE Efoetse, 17.4 km 170°S Beheloka	-24,047	43,753	40 m	Fisher-Griswold Arthropod Team	3/18/2002	2w
spinosus sp. n. HT	CASENT0443515	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n. PT	CASENT0443515	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n. PT	CASENT0443531	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0454095	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	3w
spinosus sp. n.	CASENT0454237	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0454238	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0454100	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0001365	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	1w
spinosus sp. n.	CASENT0001366	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	1w
spinosus sp. n.	CASENT0003947	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0001369	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0454236	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0057339	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	B.L.Fisher	4/16/2005	1w
spinosus sp. n.	CASENT0454094	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0443504	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0443512	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0443601	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0442542	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0443593	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0443501	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0443502	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0442540	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0442541	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	1w
spinosus sp. n.	CASENT0443539	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0443544	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0443540	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	2w
spinosus sp. n.	CASENT0001469	Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61°ENE Tsimelahy, 36.1 km 308°NW Tolagnaro	-24,93	46,646	300 m	Fisher-Griswold Arthropod Team	1/16/2002	2w
spinosus sp. n.	CASENT0443605	Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325°NW Amboasary	-24,930	46,210	65 m	Fisher-Griswold Arthropod Team	2/7/2002	3w
spinosus sp. n.	CASENT0108875	Anosy Region, Distric of Amboasary,58Km SW of Fort Dauphin, 08Km NW of Amboasary, Berenty Special Reserve	-25,021	46,306	36 m	Mike, Rin’ha	11/30/2003	1w

Figure 13.

Dendrogram for NC-clustering scores with AU/BP values (%), classification of objects based on recursive partitioning with mesosomal profile of four species of hafahafa species-group is mapped on distributional map of Madagascar. Abbreviations: AU = approximately unbiased P-value, BP = bootstrap probabilities before statistical adjustments. Rectangles show the final species hypothesis. Color codes: Nesomyrmex capricornis sp. n. (yellow), Nesomyrmex hafahafa sp. n. (red), Nesomyrmex medusus sp. n. (blue), Nesomyrmex spinosus sp. n. (green).