Taxonomic revision of the Malagasy Camponotus grandidieri and niveosetosus species groups (Hymenoptera, Formicidae) using qualitative and quantitative morphology
Author
Rakotonirina, Jean Claude
Author
Csősz, Sándor
Author
Fisher, Brian L.
text
Zootaxa
2017
2017-03-03
4238
2
203
245
journal article
36356
10.11646/zootaxa.4238.2.2
0a267faa-8b7b-44e9-abf3-fb0902f6a0d7
1175-5326
345233
1030F6ED-734A-44C1-9D9C-BC94BCB7A6CD
Camponotus voeltzkowii
Forel
(
Figures 7
B, 18, 26)
Camponotus voeltzkowii
Forel, 1894
: 226
.
Lectotype
minor worker,
present designation
,
Madagascar
,
Mahajanga
(Voeltzkow),
AntWeb
CASENT0101376 (
MHNG
) [examined]
.
Paralectotype
minor worker, same data as lectotype but with specimen code CASENT0101377 (
MHNG
) [examined]. [Combination in
Camponotus (Myrmoturba)
:
Forel, 1914
: 271
. Combination in
Camponotus (Myrmopiromis)
:
Wheeler, 1922a
: 1052
;
Emery 1925
: 129
;
Bolton, 1995
: 129
, 131].
Diagnosis.
Integument black; posterior portion of head, dorsum of mesosoma, and lateral margin of propodeal declivity with randomly scattered, whitish, erect hairs; basal half of antennal scape flattened dorsoventrally; in lateral view, mesosoma short and high, propodeal dorsum shorter than declivitous face; few scattered erect hairs present immediately behind lateral margin of clypeus; dorsum of body with numerous, randomly scattered, long, erect hairs and abundant long pubescence; tibiae with sparse suberect hairs.
Description. Minor worker.
In full-face view head slightly longer than wide (CWb/CL: 0.91±0.03, 0.87– 1.07), lateral borders straight and diverging posteriorly and joining the more or less posterior margin in a distinct angle. In lateral view, posterior cephalic margin extending backward into a rounded lobe. Eyes protruding, almost breaking lateral borders of head (CWb/CL: 0.91±0.03, 0.87–1.07; CW/CL: 0.89±0.03, 0.86–1.04), located at posterior sixth to fifth portion of head (PoOc/CL: 0.17±0.01, 0.15–0.23). With head in full-face view, clypeus trapezoidal; anteromedian margin projecting into a triangular lobe; anterolateral portion bluntly angulate; median portion convex, with visible longitudinal carina. Mandible triangular, apical margin equipped with six teeth. Antennal scape long, roughly distal portion extending beyond posterior margin of head (SL/CS: 1.10±0.05, 1.01– 1.19); basal portion cylindrical, not flattened dorsoventrally. In lateral view, mesosoma short and high (MPH/ML: 0.54±0.05, 0.46–0.71), dorsal outline of promesonotum domelike, its level higher than that of propodeum; anterodorsal portion of pronotum angulate; propodeal dorsum short, joining the sharply inclined propodeal declivity into a blunt angle; metanotal groove obsolete. Opening of propodeal spiracle circular or elongate to slitlike. In lateral view, petiolar node scalelike and high, petiolar spiracle located at level lower than midheight of node; anterior face starting to incline posteriorly at dorsal third portion of node to reach the straight posterior face dorsally. Middle and hind legs with single pectinate tibial spur. Transverse light strip on posterior margin of gastral tergites wide, maximum width roughly 1/8 the visible width of second tergite.
Head dorsum finely and densely reticulate punctate to finely areolate, becoming finely imbricate posteriorly; pronotum finely and densely reticulate punctate to finely imbricate, mesonotum and propodeum finely and densely reticulate rugulose. Lateral portion of head and mesosoma finely imbricate. Gastral tergites finely and transversely strigulate anteriorly and finely imbricate posteriorly. Mandible with coriarious ponctulate sculpture. Whitish erect hairs randomly scattered from median portion of head to near its posterior margin, where they spread transversally to near the lateral margin and down anteriorly to near base of mandible and behind lateral margin of clypeus. Head, pronotum, anterior section of mesonotum, and posterior section of gastral tergites covered with hairs thinner and more erect than posterior portion of mesonotum, propodeum, petiolar node, and first gastral tergite; tibiae covered with scattered suberect hairs. Pubescence long and quite dense, distance between them smaller than their length; pubescence more abundant on gaster than mesosoma and head.
Major worker.
With characteristics of minor worker, but head subquadrate (CWb/CL: 0.97±0.02, 0.94–0.99) to cordate in full-face view, posterior cephalic margin slightly broadly excised. Eyes placed more interior relative to head capsule (CWb/CL: 0.97±0.02, 0.94–0.99; CW/CL: 0.84±0.01, 0.81–0.85). Mandibles more robust, masticatory margin with more than six teeth. Clypeus with elongate median portion (ClyL/GPD: 0.78±0.02, 0.73– 0.81), anteromedian margin straight to slightly broadly convex. Antennal scape short, barely extending beyond posterior cephalic border (SL/CS: 0.76±0.01, 0.75–0.79). Dorsal outline of mesosoma forming a continuous convexity; metanotal groove a slightly impressed line. Lateral portion of head finely and densely reticulate punctate to finely areolate, with scattered groups of two to seven areoles surrounding the base of an appressed or erect hair; each group of these areoles is sometimes embedded in a poorly defined puncture. Lateral portion of mesosoma finely areolate to finely imbricate.
Discussion.
See discussion under
C. mita
.
For
C. voeltzkowii
, the conclusions drawn from qualitative morphology-based taxonomy agree with the classification hypothesis provided by the exploratory data analysis and the confirmatory technique of the multivariate morphometrics. The combination of this information strengthens the status of
C. voeltzkowii
as a species.
FIGURE 18.
Camponotus voeltzkowii
minor worker CASENT0121619. A: lateral view. B: head in full-face view. C: dorsal view.
Distribution and biology.
Camponotus voeltzkowii
is one of the most common species within the
niveosetosus
species group of the Malagasy region. In
Madagascar
, it is widespread in the west but absent in the northernmost region. To the north, its geographic distribution is limited to montane rainforest of the northern highland. Along the western portion of the island, the distributional range of
C. voeltzkowii
consists of the dry forest on Tsingy, gallery forest, littoral forest habitats in the north to southwest, as well as spiny bush and thicket in the extreme south of the island. The species is also known to occur in Uapaca woodland of the central high plateau region, and in the northern-and southern-central portions of the island’s eastern rainforests. The species also can occupy anthropogenic habitats such as disturbed and degraded forests and urban gardens. Colony nests are found either on the forest floor under stones or in rotten logs, or in the dead branches or twigs of trees. Foraging activity is carried out on the ground and through leaf litter or on lower vegetation.
FIGURES 19–26.
Distribution maps of the
Camponotus grandidieri
and
niveosetosus
species groups in the Malagasy region. Fig. 19:
C. auropubens
;
Fig. 20:
C. efitra
;
Fig. 21:
C. maintikibo
;
Fig. 22:
C. grandidieri
;
Fig. 23:
C. descarpentriesi
, Fig. 24:
C. madagascarensis
, Fig. 25:
C. mita
, Fig. 26:
C. voeltzkowii
.
Camponotus voeltzkowii
is sympatric with
C. madagascarensis
and
C. grandidieri
.
Additional material examined. MADAGASCAR
:
Madagascar
(
MHNG
)
;
Province
Antananarivo
:
Analamanga Region
,
District
of
Ankazobe
,
Ambohitantely
,
46 km
NE of Ankazobe
, -18.198, 47.2815,
701 m
, sclerophyll forest, (
Rin'ha
,
Mike
) (
CASC
)
;
Province
Fianarantsoa
:
Fitovavy Fitovinany Region
,
District
of
Ifanadiana
,
12 km
W of Ranomafana
, -21.25083, 47.40717,
1127 m
, forest edge, open area, (
Rin'ha
,
Mike
) (
CASC
)
;
Parc National d'Isalo
,
Ambovo Springs
,
29.3 km
4° N Ranohira
, -22.29833, 45.35167,
990 m
,
Uapaca
woodland, (
Fisher
,
Griswold
et al.
) (
CASC
)
;
Parc National d'Isalo
,
Sahanafa River
,
29.2 km
351° N Ranohira
, - 22.31333, 45.29167,
500 m
, gallery forest, (
Fisher
,
Griswold
et al.
) (
CASC
)
;
radio tower,
Ranomafana National Park
,
Fianarantsoa
Prov., -21.25083, 47.40717,
1130 m
, forest edge, mixed tropical forest, open area, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
stream area,
900 m
E of
Isalo National Park Interpretive Center
,
Fianarantsoa
Prov., - 22.62667, 45.35817,
750 m
, open area near stream, (R. Harin'Hala) (
CASC
)
;
Province
Mahajanga
: [
Majanga
] (
MHNG
)
;
Ambolomaiky
,
48.1 km
SE
Mahajanga
, -15.8541, 46.74663,
77 m
, disturbed dry forest, (
B.L. Fisher
et al.
) (
CASC
)
;
Ampijoroa National Park
,
160 km
N Maevatanana
,
Mahajanga
Prov., deciduous forest, -16.31944, 46.81333,
43 m
, deciduous forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Boeny Region
,
District
of
Soalala
,
Beaboaly Bamboo forest
,
10 km
SW of Soalala
,
4 km
from
Baly village
, -16.04533, 48.804, 9 m, bamboo forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Boeny Region
,
District
of
Marovoay
,
Ampijoroa National Park
,
160 km
N of Maevatanana
on RN 0 4, -16.31933, 46.81333,
42 m
, deciduous forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Boeny Region
,
District
of
Soalala Analamanitra forest
,
14 km
SW of Mitsinjo
, -16.7, 45.7,
19 m
, dense dry forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Forêt de Tsimembo
,
8.7 km
336° NNW Soatana
, -19.02139, 44.44067,
20 m
, tropical dry forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Parc National Tsingy de Bemaraha
,
2.5 km
62° ENE Bekopaka
,
Ankidrodroa River
, -19.13222, 44.81467,
100 m
, tropical dry forest on
Tsingy
, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Parc National Tsingy de Bemaraha
,
3.4 km
93° E Bekopaka
,
Tombeau Vazimba
, - 19.14194, 44.828, 50 m, tropical dry forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Region Sofia
,
Bemanevika
, -14.337, 48.58874,
1606 m
, montane rainforest, (
B.L. Fisher
et al.
) (
CASC
)
;
Réserve
forestière
Beanka
,
50.7 km
E Maintirano
, -17.88021, 44.46877,
140 m
, tropical dry forest on tsingy, (
B.L. Fisher
et al.
) (
CASC
)
;
Sofia Region
,
District
of
Sofia
,
Anjiamangirana
45 km
S Antsohihy
,
Analagnambe
galleryforest,
5 km
W Anjiamangirana
, -15.157, 47.73417,
97 m
, low degraded dry forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Station Forestiere Ampijoroa
, -16.31667, 46.81667,
80 m
, tropical dry forest, (
P.S. Ward
) (
PSWC
)
;
Province
Toamasina
:
Parc National de Zahamena
,
Tetezambatana forest
, near junction of
Nosivola
and
Manakambahiny Rivers
, - 17.74298, 48.72936,
860 m
, rainforest, (
B.L. Fisher
et al.
) (
CASC
)
;
Province
Toliara
: [
Fort Dauphin
;
Museum Paris Collection
,
Ernest André
1914] (
MNHN
)
;
18.4 km
N
Amboasary
, -24.87367, 46.39767,
85 m
, spiny forest/ thicket, (
B.L. Fisher
et al.
) (
CASC
)
;
2 km
SW Mahamavo
,
Rés.Andohahela
, -24.78333, 46.7,
320 m
, tropical dry forest, (
P.S. Ward
) (
PSWC
)
;
45 km
NE
Morondava
, -20.05, 44.61667,
30 m
, tropical dry forest, (
P.S. Ward
) (
PSWC
)
;
55 km
NE
Morondava
, -19.9, 44.6,
25 m
, tropical dry forest, (
P.S. Ward
) (
PSWC
)
;
Ambohimahavelona village
33 km
NE of Tulear
,
Andoharano
dry forest, -23.44083, 43.89967,
46 m
, dry forest, (
M. Irwin
, R. Harin'Hala) (
CASC
)
;
Androy Region
,
District
of
Tsihombe
,
74 km
S of Tsihombe
,
Cap Ste Marie Reserve
, - 25.58767, 45.163, 36 m, spiny bush, (
Rin'ha
,
Mike
) (
CASC
)
;
Anosy Region
,
District
of
Amboasary
,
58 km
SW of Fort Dauphin
,
8 km
NW of Amboasary
,
Berenty Special Reserve
, -25.00667, 46.30333,
85 m
, galleryforest, (
Rin'ha
,
Mike
) (
CASC
)
;
Anosy Region
,
District
of
Amboasary
,
Andohaela National Park Parcelle
III,
Ihazofotsy
,
32 km
NE Amboasary
, -24.83083, 46.53617,
58 m
, dry forest, spiny forest, (
M.E. Irwin
,
F.D. Parker
, R. Harin'Hala) (
CASC
)
;
Anosy Region
,
District
of
Amboasary
,
58 km
SW of Fort Dauphin
,
8 km
NW of Amboasary
,
Berenty Special Reserve
, -25.021, 46.3055,
36 m
, spiny forest, (
Rin'ha
,
Mike
) (
CASC
)
;
Anosy Region
,
District
of
Fort-Dauphin
,
Andohaela National Park Parcelle II
,
Tsimela
,
42 km
W of Fort-Dauphin
, -24.93683, 46.62667,
177 m
, transition forest, (
M.E. Irwin
,
F.D. Parker
, R. Harin'Hala) (
CASC
)
;
Anosy Region
,
Parc National d'Andohahela
,
Forêt de Manatalinjo
, -24.82505, 46.57811,
90 m
, spiny forest/thicket, (
B.L. Fisher
,
F.A. Esteves
et al.
) (
CASC
)
;
Anosy Region
,
Parc National d'Andohahela
,
Forêt de Manatalinjo
, -24.82466, 46.60111,
100 m
, spiny forest/ thicket, (
B.L. Fisher
,
F.A. Esteves
et al.
) (
CASC
)
;
Berenty Reserve
, -24.98333, 46.3,
30 m
, gallery forest, (
D.M. Olson
) (
PSWC
)
.
,
Berenty Special Reserve
,
8 km
NW Amboasary
,
58 km
SW of Fort Dauphin
,
Tulear Province
, - 25.00667, 46.30333,
85 m
, gallery forest, (
M.E. Irwin
,
F.D. Parker
, R. Harin'Hala) (
CASC
)
;
Ejeda
, -24.3505, 44.516, 250 m, urban/garden, (
B.L. Fisher
et al.
) (
CASC
)
;
Forêt de Kirindy
,
15.5 km
64° ENE Marofandilia
, - 20.06915, 44.66041667,
30 m
, tropical dry forest, (
B.L. Fisher
) (
CASC
)
;
Forêt de Kirindy
,
15.5 km
64° ENE Marofandilia
, -20.045, 44.66222,
100 m
, tropical dry forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Forêt de Kirindy
,
15.5 km
64° ENE Marofandilia
, -20.045, 44.66222,
100 m
, tropical dry forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Forêt de Mahavelo
,
Isantoria River
, -24.75833, 46.15717,
110 m
, spiny forest/thicket, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Forêt de Mahavelo
,
Isantoria River
,
5.5 km
37° NE Ifotaka
, - 24.75361, 46.1515,
115 m
, spiny forest/thicket, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Forêt de Mite
,
20.7 km
29° WNW Tongobory
, -23.52417, 44.12133,
75 m
, gallery forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Forêt Vohidava
88.9 km
N Amboasary
, -24.24067, 46.28783,
500 m
, spiny forest/dry forest transition, (
B.L. Fisher
et al.
) (
CASC
)
;
Forêt Vohidava
89.2 km
N Amboasary
, -24.239, 46.28233,
850 m
, tropical dry forest, (
B.L. Fisher
et al.
) (
CASC
)
;
Forêt Vohidava
89.2 km
N Amboasary
, -24.239, 46.28233,
850 m
, tropical dry forest, (
B.L. Fisher
et al.
) (
CASC
)
;
Forêt Vohidava
89.6 km
N Amboasary
, -24.23333, 46.30167,
230 m
, spiny forest/thicket, (
B.L. Fisher
et al.
) (
CASC
)
;
Ihazofotsy - Parcel
III,
Andohahela National Park
, transition forest,
Tulear Province
, - 24.83483, 46.48683,
80 m
, transition between spiny and dry deciduous forests, (
M.E. Irwin
,
F.D. Parker
, R. Harin'Hala) (
CASC
)
;
Kirindy
,
47.3 km
NE Morondava
, -20.07125, 44.6655,
60 m
, tropical dry forest, (
B.L. Fisher
et al.
) (
CASC
)
;
Makay Mts.
, -21.227, 45.33222,
475 m
, gallery forest on sandy soil, (
B.L. Fisher
et al.
) (
CASC
)
;
Makay Mts.
, -21.21985, 45.32396,
500 m
, gallery forest on sandy soil, (
B.L. Fisher
et al.
) (
CASC
)
;
Makay Mts.
, - 21.23343, 45.32913,
460 m
, gallery forest with bamboo, (
B.L. Fisher
et al.
) (
CASC
)
;
Manombo
, -22.80707, 43.76375,
222 m
, gallery forest, TS1, (
Frontier Wilderness Project
) (
CASC
)
;
Menabe Region
,
District
of
Morondava
,
Beroboka village
45 km
NE of Morondava
,
Antsarongaza
, dry forest
7.5 km
E of Beroboka
, -19.9775, 44.66633,
50 m
, dry forest, (
Michael Irwin
,
Rin'ha
) (
CASC
)
;
Menabe Region
,
District
of
Morondava
,
Beroboka village
,
45km
NE of Morondava
,
Antsarongaza
, gallery forest,
7 km
E of Beroboka
, -19.9775, 44.66533,
45 m
, gallery forest, (
Michael Irwin
,
Rin'ha
) (
CASC
)
;
Parc National d'Andohahela
,
Forêt d'Ambohibory
,
1.7 km
61° ENE Tsimelahy
,
36.1 km
308° NW Tolagnaro
, -24.93, 46.6455,
300 m
, tropical dry forest, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Parc National d'Andohahela
,
Forêt de Manatalinjo
,
33.6 km
63° ENE Amboasary
,
7.6 km
99° E Hazofotsy
, -24.81694, 46.61,
150 m
, spiny forest/thicket, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Parc National de Kirindy Mite
,
16.3 km
127° SE Belo sur Mer
, -20.79528, 44.147, 80 m, tropical dry forest, (Fisher-
Griswold Arthropod Team
) (
CASC
)
;
Parc National de Tsimanampetsotsa
,
Forêt de Bemanateza
,
20.7 km
81° E Efoetse
, 23.0 km 131°
SE Beheloka
, -23.99222, 43.88067,
90 m
, spiny forest/thicket, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
Parcel I
,
Beza Mahafaly Reserve
, near research station,
Tulear Province
, -23.6865, 44.591, 165 m, dry deciduous forest, (R. Harin'Hala) (
CASC
)
;
Parcel II
,
Beza Mahafaly Reserve
, near
Bellevue
,
Tulear Province
, - 23.68983, 44.5755,
180 m
, spiny forest, (R. Harin'Hala) (
CASC
)
;
Res. Beza Mahafaly,
Parcel
1, -23.65, 44.63333,
130 m
, tropical dry forest, (
P.S. Ward
) (
PSWC
)
;
Rés. Beza Mahafaly,
Parcel
2, -23.68333, 44.56667,
200 m
, spiny forest/thicket, (
P.S. Ward
) (
PSWC
)
;
Réserve Privé Berenty
,
Forêt d'Anjapolo
,
21.4 km
325° NW Amboasary
, - 24.92972, 46.20967,
65 m
, spiny forest/thicket, (
Fisher-Griswold Arthropod Team
) (
CASC
)
;
southern Isoky-
Vohimena Forest
,
59 km
NE Sakaraha
, -22.46667, 44.85,
730 m
, tropical dry forest, (
B.L. Fisher
) (
CASC
)
.