The genus Camponotus Mayr, 1861 is one of the most speciose and widespread genera within the Formicidae, comprising more than 90 described species and subspecies in the Malagasy region. This genus has received renewed attention over the last five years. In 2016, the Camponotus edmondi species group was revised by Rakotonirina et al. (2016). Later, Camponotus grandidieri and niveosetosus species groups, the subgenera Myrmopytia and Mayria have been revised (Rakotonirina et al. 2017;Rasoamanana et al. 2017;Rakotonirina and Fisher 2018). In total, 41 species were included in these studies, which combined qualitative morphological analysis, multivariate morphometry, and ecological data. Here we continue to explore the taxonomy of Malagasy Camponotus and revisit the subgenus Mayria. We redefine the subgenus and describe additional species using morphological features combined with morphometry.
In the 2018 revision by Rakotonirina et al.,14 species were recognized in the subgenus. However, our further studies of Malagasy Camponotus species have revealed additional morphological traits leading to new groupings. This revision increases the size of subgenus Mayria with the transfer of 15 species from the edmondi species group (Rakotonirina et al. 2016) and six species from niveosetosus species group (Rakotonirina et al. 2017), and the description of eleven new species and the redescription of six previously described species. Together, these changes increase the total number of species in this subgenus to 39. We divide this subgenus into nine species groups: alamaina group, antsaraingy group, darwinii group, edmondi group, efitra group, ellioti group, madagascarensis group, repens group, and robustus group.
This study is based on specimens collected from the ant inventory of Madagascar project as part of a joint collaboration between CAS and Parc Botanique et Zoologique Tsimbazaza under the aegis of the Ministry of Higher Education in Madagascar. After morphological examination, all workers were separated into morphospecies, and measurements of the 19 characters were recorded for specimens in good condition. The data were subjected to a multivariate analysis, which utilized the Nest Centroid (NC)-Clustering method to determine whether differences existed among workers within the same species. The degree of inclination of pilosity is particularly important for diagnosing groups or species. In this context, we use the terms erect, suberect, subdecumbent, decumbent, and appressed following Wilson (1955).
Specimens were deposited in the following collections:
California Academy of Sciences, San Francisco, CA, USA MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland MNHN Muséum national d'Histoire naturelle, Paris, France MSNG Museo Civico di Storia Naturale di Genova "Giacomo Doria", Genova, Italy NHMB Naturhistorisches Museum, Basel, Switzerland PBZT Parc Botanique et Zoologique de Tsimbazaza, Antananarivo, Madagascar PSWC P. S. Ward Collection, University of California at Davis, CA, USA ZMHB Museum für Naturkunde, Humboldt-Universität, Berlin, Germany
All morphological observations were made with a Leica MZ9.5 stereomicroscope. We morphometrically investigated 307 individual workers belonging to 215 collecting events for the eleven newly described species and six redescribed species. Camponotus samples were collected across Madagascar by BLF and the Madagascar Biodiversity Center team (Table 1). The material is deposited in the California Academy of Sciences (CAS), San Francisco, USA. Data for all pinned specimens examined in this study are available on the web portal AntWeb (https://www.antweb.org) and can be accessed using their unique identifying specimen code (e.g., CASENT0188388). Images are linked to their specimens via the unique specimen code affixed to each pin. Type material and samples used in the morphometric analysis are given for each species after "Additional material examined" in the following order: Province, locality name, latitude, longitude, elevation (m), habitat, collector and abbreviation of depository. The most common collectors are abbreviated as follows:
BL Fisher and the Madagascar Biodiversity Center team FGAT Fisher-Griswold Arthropod Team ARA Andrianjaka Ravelomanana PSW Phil Ward
Madagascar Malaise trap program by Mike Irwin and Rasolondalao Harin'hala Hasinjaka
Digital color montage images were created using a JVC KY-F75 digital camera and syncroscopy Auto-Montage software (version 5.0), or a Leica DFC 425 camera in combination with the Leica Application Suite software (version 3.8). Distribution maps were generated by using QGIS 2.4.0 software (QGIS Development Team 2014).
Measurements were taken with a Leica MZ 9.5 stereomicroscope equipped with a cross-scaled ocular micrometer. Each worker was evaluated using 19 continuous morphometric traits, measured as in Rakotonirina et al. (2016) for the Camponotus edmondi group. The morphometric data are expressed in mm. The following measurements were used (see Figure 1):
PoOC Postocular distance. The longest distance between the posteromedian margin of the head and theposterior margins of the two compound eyes. Measured at a right angle to the reference line in full-face view (Fig. 1A). PrAC Preocular distance. The longest distance between the anteromedian margin of the clypeus and the level of the anterior margin of the compound eyes as a reference line. Measured at a right angle to the reference line in full-face view (Fig. 1A).
Scape length. Straight line length of the first antennal segment excluding the basal condyle (Fig. 1B). TCD Torular carina distance. The minimum distance between the torular arches that surround the antennal insertion (Fig. 1A).
After measuring workers (240 minors and 67 majors), multivariate statistical analysis based on the protocol described by Seifert et al. (2014) and extended by Csősz and Fisher (2016) was performed with data from minor workers only, since Camponotus workers exhibit different forms of allometry. Species hypotheses were generated using [0.81, 0.88] [0.08, 0.09] [0.19, 0.21] [0.09, 0.10] [0.23, 0.25] [0.32, 0.34] [0.30, 0.32 .12, 1.20] [0.23, 0.27] [0.54, 0.55] [0.32, 0.35] [0.83, 0.92] [0.79, 0.82] [0.84, 0.85 00, 2.33] [0.07, 0.09] [0.21, 0.23] [0.10, 0.12] [0.22, 0.28] [0.31, 0.32] [0.32, 0.35 94, 3.55] [0.26, 0.27] [0.52, 0.56] [0.16, 0.30] [0.46, 0.80] [0.77, 0.81] [0.95, 0.99] chrislaini minor (N=5) 0.95±0.17 0.12±0.03 0.18±0.01 0.11±0.02 0.26±0.03 0.36±0.03 0.34±0.03 [0.87, 1.02] [0.11, 0.13] [0.18, 0.19] [0.10, 0.12] [0.25, 0.29] [0.36, 0.38] [0.32, 0.35 [0.84, 0.93] [0.08, 0.10] [0.18, 0.20] [0.08, 0.11] [0.20, 0.26] [0.33, 0.35] [0.32, 0.34 .12, 1.24] [0.25, 0.26] [0.54, 0.54] [0.33, 0.34] [0.85, 0.91] [0.83, 0.84] [0.85, 0.87 .22, 1.63] [0.06, 0.09] [0.20, 0.23] [0.10, 0.13] [0.23, 0.28] [0.33, 0.37] [0.35, 0.41 .70, 2.47] [0.19, 0.24] [0.54, 0.59] [0.20, 0.31] [0.48, 0.81] [0.85, 0.93] [0.94, 1.20 [0.21, 0.23] [0.11, 0.13] [0.25, 0.30] [0.31, 0.34] [0.33, 0.39 [3.90, 4.33] [0.26, 0.29] [0.55, 0.57] [0.28, 0.32] [0.79, 0.92] [0.78, 0.82 [0.81, 0.86] [0.08, 0.10] [0.18, 0.20] [0.10, 0.10] [0.22, 0.25] [0.34, 0.37] [0.32, 0.34] major (N=3) 1.13±0.05 0.26±0.01 0.51±0.01 0.34±0.00 0.83±0.00 0.85±0.01 0.90±0.02 [1. 09, 1.18] [0.25, 0.27] [0.50, 0.52] [0.33, 0.34] [0.83, 0.83] [0.83, 0.86] [0.88, 0.91 [0.12, 0.13] [0.09, 0.11] [0.35, 0.38] [0.19, 0.20] [0.10, 0.12] [0.24, 0.28] [0.07, 0.08] major (N=3) 0.36±0.01 0.26±0.02 0.70±0.02 0.53±0.02 0.34±0.02 0.64±0.01 0.23±0.01 [0.35, 0.37] [0.25, 0.28] [0.69, 0.72] [0.50, 0.55] [0.32, 0.35] [0.63, 0.65] [0.22, 0.25 [0.14, 0.15] [0.09, 0.10] [0.42, 0.45] [0.26, 0.28] [0.16, 0.19] [0.22, 0.23] [0.11, 0.15] major (N=6) 0.35±0.01 0.24±0.01 0.81±0.04 0.66±0.03 0.31±0.02 0.55±0.03 0.21±0.02 [0.35, 0.36] [0.23, 0.24] [0.76, 0.88] [0.61, 0.71] [0.27, 0.34] [0.52, 0.60] [0.19, 0.25 [0.17, 0.18] [0.10, 0.10] [0.45, 0.48] [0.21, 0.23] [0.14, 0.15] [0.24, 0.29] [0.07, 0.11 [0.44, 0.47] [0.26, 0.28] [0.74, 0.83] [0.54, 0.58] [0.34, 0.36] [0.59, 0.67] [0.20, 0.23 [0.14, 0.16] [0.10, 0.12] [0.37, 0.40] [0.18, 0.19] [0.13, 0.14] [0.24, 0.27] [0.10, 0.11] major (N=2) 0.41±0.02 0.28±0.01 0.75±0.04 0.49±0.01 0.33±0.01 0.62±0.03 0.28±0.00 [0.40, 0.42] [0.27, 0.28] [0.73, 0.78] [0.49, 0.50] [0.32, 0.33] [0.60, 0.64] [0.28, 0.28 [0.13, 0.15] [0.10, 0.12] [0.37, 0.43] [0.19, 0.23] [0.11, 0.16] [0.20, 0.26] [0.09, 0.13] major (N=14) 0.35±0.01 0.27±0.01 0.83±0.08 0.56±0.02 0.30±0.03 0.59±0.03 0.25±0.03 [0.33, 0.37] [0.25, 0.28] [0.75, 0.99] [0.53, 0.58] [0.23, 0.36] [0.53, 0.67] [0.21, 0.29 [0.11, 0.12] [0.08, 0.10] [0.34, 0.46] [0.21, 0.25] [0.11, 0.14] [0.20, 0.28] [0.06, 0.12] major (N=7) 0.29±0.01 0.23±0.01 0.61±0.02 0.59±0.02 0.26±0.01 0.63±0.02 0.17±0.02 [0.27, 0.30] [0.22, 0.23] [0.58, 0.65] [0.56, 0.61] [0.24, 0.28] [0.61, 0.66] [0.14, 0.19 [0.13, 0.16] [0.10, 0.11] [0.39, 0.43] [0.18, 0.21] [0.11, 0.15] [0.23, 0.26 [0.35, 0.36] [0.25, 0.27] [0.74, 0.79] [0.50, 0.52] [0.28, 0.33] [0.63, 0.68] [0.24, 0.25 0.38, 0.40] [0.16, 0.19] [0.10, 0.11] [0.12, 0.15] --major (N=3) 0.84±0.04 0.46±0.01 0.21±0.01 0.39±0.01 --- [0.81, 0.88] [0.44, 0.46] [0.20, 0.21] [0.38, 0.40 [0.16, 0.18] [0.09, 0.11] [0.20, 0.22] --major (N=6) 0.94±0.04 0.43±0.01 0.20±0.01 0.51±0.01 --- [0.89, 0.99] [0.42, 0.44] [0.20, 0.21] [0.49, 0.53 .13, 1.30] [0.10, 0.11] [0.17, 0.20] [0.09, 0.10] [0.19, 0.22] [0.32, 0.34] [0.33, 0.36 0.91, 1.09] [0.09, 0.12] [0.19, 0.20] [0.07, 0.10] [0.16, 0.23] [0.33, 0.36] [0.31, 0.34 .61, 1.76] [0.29, 0.30] [0.49, 0.53] [0.29, 0.30] [0.81, 0.88] [0.82, 0.85] [0.89, 0.92 .31, 1.66] [0.07, 0.09] [0.21, 0.23] [0.11, 0.13] [0.23, 0.28] [0.32, 0.34] [0.35, 0.38 .43, 1.77] [0.07, 0.09] [0.21, 0.23] [0.10, 0.12] [0.23, 0.29] [0.33, 0.37] [0.35, 0.40 [2.59, 2.85] [0.23, 0.25] [0.53, 0.59] [0.23, 0.32] [0.57, 0.86] [0.82, 0.89] [1.02, 1.07] ihazofotsy minor (N=2) 0.78±0.12 0.09±0.03 0.20±0.04 0.10±0.02 0.25±0.07 0.31±0.00 0.34±0.01 [0.75, 0.81] [0.08, 0.10] [0.19, 0.21] [0.10, 0.11] [0.23, 0.27] [0.31, 0.31] [0.34, 0.35 .53, 1.98] [0.07, 0.08] [0.22, 0.23] [0.10, 0.12] [0.25, 0.30] [0.31, 0.34] [0.34, 0.38 [2.26, 3.15] [0.22, 0.25] [0.55, 0.58] [0.28, 0.34] [0.73, 0.93] [0.82, 0.87] [0.99, 1.11 0.77, 0.81] [0.08, 0.09] [0.19, 0.19] [0.09, 0.10] [0.21, 0.25] [0.31, 0.34] [0.28, 0.29 0.15, 0.16] [0.11, 0.12] [0.38, 0.40] [0.19, 0.22] [0.14, 0.15] [0.25, 0.28] [0.10, 0.11 0.16, 0.18] [0.12, 0.14] [0.32, 0.34] [0.18, 0.20] [0.11, 0.14] [0.24, 0.26 0.14, 0.15] [0.11, 0.13] [0.37, 0.42] [0.21, 0.23] [0.11, 0.13] [0.22, 0.24] [0.10, 0.13 0.13, 0.15] [0.10, 0.12] [0.33, 0.41] [0.20, 0.24] [0.12, 0.17] [0.21, 0.28] [0.10, 0.14 0.33, 0.35] [0.25, 0.28] [0.68, 0.91] [0.52, 0.57] [0.30, 0.39] [0.54, 0.70] [0.21, 0.34 0.13, 0.15] [0.10, 0.11] [0.37, 0.41] [0.20, 0.23] [0.11, 0.14] [0.22, 0.26] [0.10, 0.13 0.33, 0.33] [0.24, 0.27] [0.72, 0.79] [0.52, 0.58] [0.31, 0.33] [0.61, 0.69] [0.25, 0.29 [0.12, 0.14] [0.09, 0.10] [0.38, 0.44] [0.20, 0.22] [0.13, 0.15] [0.22, 0.26] [0.11, 0.15 [0.19, 0.34] [0.23, 0.27] [0.71, 0.79] [0.52, 0.56] [0.30, 0.36] [0.59, 0.70] [0.22, 0.28 0.13, 0.15] [0.09, 0.10] [0.40, 0.42] [0.17, 0.20] [0.12, 0.13] [0.26, 0.27 0.98, 1.20] [0.49, 0.54] [0.19, 0.22] [0.44, 0.46 0.37, 0.38] [0.16, 0.17] [0.11, 0.11] [0.10, 0. 0.92, 1.14] [0.08, 0.09] [0.22, 0.23] [0.11, 0.11] [0.24, 0.24] [0.38, 0.38] [0.39, 0.40 .71, 1.77] [0.25, 0.26] [0.53, 0.54] [0.26, 0.31] [0.70, 0.77] [0.90, 0.93] [0.97, 1.00 0.13, 0.15] [0.11, 0.12] [0.37, 0.39] [0.20, 0.22] [0.13, 0.13] [0.21, 0.29] [0.11, 0.13 0.34, 0.40] [0.26, 0.30] [0.78, 0.86] [0.54, 0.58] [0.27, 0.37] [0.56, 0.64] [0.26, 0.33 0.15, 0.16] [0.12, 0.12] [0.33, 0.36] [0.21, 0.22] [0.13, 0.14] [0.24, 0.25] [0.08, 0.11] major (N=2) 0.38±0.00 0.31±0.01 0.66±0.02 0.54±0.00 0.27±0.00 0.64±0.02 0.23±0.01 [0.38, 0.38] [0.30, 0.31] [0.65, 0.68] [0.54, 0.54] [0.27, 0.28] [0.63, 0.65] [0.22, 0.23 0.78, 0.80] [0.44, 0.46] [0.21, 0.22] [0.42, 0.42] ---
the NC-clustering technique (Seifert et al. 2014) via the packages CLUSTER (Maechler et al. 2014) and MASS (Venables and Ripley 2002). The procedure followed Csősz and Fisher (2016) and Rakotonirina et al. (2017Rakotonirina et al. ( , 2018)).
The subgenus Mayria Forel Mayria Forel, 1878: 369. [As subgenus of Camponotus Forel, 1894: 227;Bolton, 1994: 50]. Camponotus Forel, 1894e: 227;Forel, 1914a: 262;Forel, 1927: 251;Wheeler, 1922: 706;Emery, 1925dEmery, : 1121;;subsequent authors: Bolton, 1995b: 34;Bolton, 2003: 113. Mayria as junior synonym of Myrmosaga Forel, 1912. Myrmepinotus Santschi, 1921: 312. [As subgenus of Camponotus]. Type species: Camponotus echinoploides Forel, 189. Syn. nov.
Diagnosis. Workers of the subgenus Mayria are easily distinguished from the other subgenera in Madagascar by the following: postocular distance shorter (PoOC/CL< 0.12); clypeus with short median lobe, its anterolateral corner rounded; anterior margin of pronotum strongly convex in lateral view, forming a rounded flange and extended laterally to form an obtuse humeral angle (Fig. 2A-C); in dorsal view, pronotal disc rectangular with distinct lateral margin (Fig. 2D-F); propodeum dorsum never concave; posterior margin of petiolar node relatively sharp and always surmounted with standing hairs. Body integument opaque, sculptured, and generally matte in color.
Standing setae consist of straplike hairs, at least thickened basally (Fig. 2F,G).
Diagnosis of minor worker. Head of minor trapezoidal, broadened posteriorly, lateral cephalic margins slightly convex to straight, occipital margin almost straight to evenly convex (Fig. 2H-J). Mandible short and triangular, armed with six teeth, basal margin forming a smooth curve, not forming a right angle with the mandible insertion. Palp formula: 6, 4. Maxillary palp long. Clypeus transversely trapezoidal with rounded to evenly triangular or rectangular anterior margin, anterolateral portion always rounded, posterior margin of clypeus straight. Antennae 12-segmented: moderately long, scape gradually broadened apically. Frontal lobe narrowed anteriorly, wider behind the antennal insertion; frontal carina V-shaped. Compound eye moderately sized, protruding, breaking the outline of the lateral cephalic margin in alamaina species group, located posterior to the midlength of the head.
Mesosoma in lateral view, dorsal outline of mesosoma continuous (darwinii group) to complex (other groups). Promesonotal suture visible. Pronotum: humeral angle tuberculate (C. ellioti species group), marginate in the remaining species group; in dorsal view, pronotal disc usually rectangular. Mesonotum in dorsal view, wider than long; laterally marginate in C. edmondi species group. Mesopleuron and propodeal surface together distinctly longer than lateral portion of pronotum in lateral view. Propodeal lobe reduced. Metapleural gland absent. Procoxa of normal to moderate size for C. edmondi species group. Middle and hind tibiae with single pectinate spur.
Petiolar node: broad and massive, ranging from squamiform to nodiform, posterodorsal edge of petiole always marginate and ornamented with thick, erect hairs.
Integument opaque and sculptured, ranging from finely and densely reticulatepunctate to finely and densely imbricate.
Pilosity: standing setae thick at least at the base, whitish to pale colored, erect to appressed hairs, filiform to spatulate hairs, moderately distributed on entire dorsum of head and body; ground pilosity always present and conspicuous.
Diagnosis of major worker. Most of the morphological traits cited above for minor workers are also characteristics of major workers. However, major workers exhibit the following characters: head more subquadrate to rectangular; malar area thimblelike with weak impression at the base of each seta (Fig. 2K-M); lateral portion of head sculptured as finely and densely reticulate-punctate, imbricate, or areolate, and superimposed with two to seven smaller areoles embedded in scattered large punctures, from which an appressed hair arises medially; antennal scape shorter, its apex barely surpassing or not extending beyond posterior cephalic margin; dorsal outline of mesosoma with the same structure as minor worker.
Note. The type species Camponotus echinoploides of the subgenus Myrmepinotus shares the characters of the Mayria diagnosis and therefore Myrmepinotus is synonymized with Mayria. The following characters place C. echinoploides within the Mayria subgenus: in dorsal view, mesosoma relatively short and distinctly marginate anteriorly and laterally; pronotum and mesonotum widened; promesonotal and metanotal sutures well marked; in profile, petiolar node thick with convex anterior and posterior margins, and body covered with whitish, thick hairs. In fact, the lectotype of C. repens and the holotype of C. echinoploides share a suite of morphological characters: in fullface view, head elongate, broadened posteriorly with slightly convex sides; clypeus with short median lobe; in profile, anterior margin of pronotum strongly convex; pronotal disc subrectangular, marginate laterally; petiolar node as high as long; standing setae consist of thick, whitish hairs; body integument dull. These morphological characters are observed also in specimens belonging to the edmondi species group but absent for species that are removed from the subgenus Mayria (Table 2). Accordingly, we have treated Myrmepinotus as a synonym of Mayria.
Minor worker: in full-face view, head subovate, distinctly longer than broad, lateral margin more or less straight and rounding to the broadly convex posterior margin (Fig. 3A). Anteromedian margin of clypeus generally convex; posteromedian margin slightly notched.
Major worker: in full-face view, head subquadrate, posterior margin approximately straight and rounding to lateral margins. Clypeus hexagonal, its anterior border straight and medially excised, frontal triangle distinct (Fig. 3C).
Minor and major workers: Mesosomal profile almost flattened and interrupted by the impressed metanotal suture. Promesonotal dorsum flattened; anterodorsal angle of pronotum and dorsolateral portion of mesosoma bluntly marginate; posterolateral margin of propodeum rounding to declivitous surface (Fig. 3B-D). In dorsal view, mesonotum as long as broad, but sides converging posteriorly; mesonotum and propodeum laterally compressed at their junction; metanotal groove vestigial, represented by a transverse line. In profile, petiolar node anteroposteriorly flattened and tapered dorsally; anterior margin slightly convex and posterior margin more or less straight; dorsal margin straight or weakly excised medially (Fig. 3D). Dorsum of head, mesosoma, and petiole with imbricate sculpture; gaster with finer imbrication; mandible coriariouspuncticulate. Pairs of erect hairs transversely arranged on mesosomal dorsum and in a row on gastral tergite, pubescence short and scattered on dorsum of body.
Remarks. The group is distinguishable from all other members of the subgenus Mayria by the combination of the following characters: mesosoma distinctly marked with promesonotal and metanotal sutures; petiole with sharp edge, its anterior margin convex and its posterior margin more or less straight; propodeal spiracle placed anterior to posterolateral margin of propodeum; head and mesosoma black to dark brown, gaster and appendages dark brown to yellow or depigmented yellow; cervical shield joining pronotal dorsum directly; junction of dorsal face to lateral face of propodeum without sharp carina, posterolateral margin present.
Even though Rakotonirina et al. (2016) described these three species as members of C. edmondi species group, these character states justify a different grouping. In addition, species of this group prefer arid habitats such as dry forest, spiny bush and thicket, and gallery forest, with the exception of Camponotus bevohitra, which can be found in montane rainforest.
Minor worker: in full-face view, head subtriangular, longer than wide, with arched posterior margin and straight lateral sides. Clypeus with projecting triangular lobe (Fig. 4A).
Major worker: in full-face view, head large, about as broad as long, with slightly convex posterior and lateral borders. Anterior margin of clypeus forming a rounded lobe (Fig. 4C).
Minor and major workers: antennal scape entirely flattened longitudinally. In lateral view, mesosoma short and high, propodeal dorsum relatively shorter than declivi-tous face. Suberect spatulate hairs present on occipital portion until anterior margin of eyes and on clypeal dorsum, pubescence long and distinct, moderately abundant next to antennal insertion. Body massive, black throughout. In lateral view, petiole cuneiform, with sharp dorsal margin. Body integument matte black, finely and densely reticulate-punctate. Dorsum of mesosoma and gaster covered with numerous spatulate, whitish hairs (Fig. 4B,D).
Remarks. Camponotus antsaraingy is placed in its own group, based mainly on the following characters: its large size, the shape of its antennal scape, propodeum distinctly reduced, its dorsum covered with erect spatulate setae, and having only three gastral tergites visible in dorsal view. It is one of the groups in the subgenus Mayria with white pilosity. It differs from the madagascarensis species group by the structure of its mesosoma and its larger size.
Minor worker: in full-face view, head more or less rectangular, with convex lateral sides and evenly straight occipital margin. Setae on dorsum of head sparsely distributed on occipital portion, arranged in longitudinal row along frontal carina (Fig. 5A). Major worker: in full-face view, head somewhat longer than broad, a little narrower in front than broad (Fig. 5C).
Minor and major worker: clypeus with short, truncated, or rounded rectangular median lobe, its dorsum covered with randomly spaced setae. Mesosomal profile continuous, promesonotal and metanotal suture not impressed, sometimes forming a shiny line. Petiole scale usually not thick, but thin or sharp along margins. Integument opaque, matte black or brown, distinctly reticulate-punctate throughout. Body mostly covered with coarse yellow, red, or white setae, sometimes forming a dense fur coating the gaster or mesosoma (Fig. 5B,D).
Remarks. Camponotus darwinii species group has consistently been considered a distinctive group because of the presence of coarser and denser pubescence on its dorsal margin; integument dull and sculptured; dorsal outline of mesosoma continuous. This group is distinguishable from all other members of the subgenus Mayria by the presence of dense, whitish pilosity on its dorsum and the arcuate profile of its mesosoma. Species in this group are traditionally classified under subgenus Myrmopiromis, type species Camponotus fulvopilosus, type location South Africa. The subgeneric definition by Emery (1925) and the revised definition of Camponotus fulvopilosus species group by Robertson (1990Robertson ( , 1997) ) are quite different from Camponotus darwinii species group. In addition to the species distribution, the number of mandibular teeth is a good character for distinguishing these two species groups: six for C. darwinii group and seven or eight for C. fulvopilosus group.
Minor worker: in full-face view, head as long as broad, posterior margin broadly convex, lateral margins roughly straight (Fig. 6A).
Major worker: in full-face view, head subquadrate, posterior and lateral margins more or less straight to convex (Fig. 6C).
Minor and major: clypeus with broadly convex anterior margin and medially notched posterior margin. In profile, dorsal outline of mesosoma interrupted by metanotal suture, promesonotal suture distinct but not impressed. In profile, petiole always biconvex. Body integument opaque, densely reticulate-punctate (Fig. 6B-D). Few scattered hairs present on mesosomal dorsum, pubescence abundant to reduced.
Remarks. The Camponotus edmondi species group can be distinguished by the combination of the following characters: dorsolateral margin of propodeum marginate or extending into a sharp ridge, propodeal declivity usually concave, anterolateral corner of pronotum most often marginate, forecoxa larger than the width of mesopleuron, and propodeal dorsum abruptly sloping down to the insertion of the petiole.
Minor worker: in full-face view, head subovate with strongly convex posterior and lateral sides. Anterior clypeal margin broadly rounded (Fig. 7A).
Major worker: in full-face view, head roughly quadrate or rectangular, head sides evenly convex (as in C. efitra) to straight (as in C. chrislaini) (Fig. 7C).
Minor and major: in profile, metanotal suture impressed, the propodeal dorsum forming a separate convexity from a domelike promesonotum, dorsum of propodeum below level of promesonotum. Clypeus evenly convex to straight in profile, its anterior margin convex to broadly triangular, not projecting, posterior margin weakly notched medially, major worker with massive mandible, occipital portion asetose, few standing setae present between frontal carinae but moderately present on clypeal dorsum and malar area. Petiole nodiform. Color brown. Integument sculpture finely imbricate to glabrous, apparently shiny (Fig. 7B,D).
Remarks. The efitra group is characterized by the color and sculpture of its integument; and a convex promesonotum separated from the propodeum by a weak metanotal suture. In Rakotonirina, Csősz & Fisher, 2017, C. efitra was considered a member of the grandidieri species group. Here, this species is separated from this group by the form of the mesosoma and the petiolar scale, the type of integument, and the general sculpture of major workers.
Minor worker: in full-face view, head subtriangular, posterior margin evenly convex, head sides anterior to eyes subparallel and diverging posteriorly (Fig. 8A).
Major worker: in full-face view, head longer than broad, subtriangular to rectangular; with weakly convex sides (Fig. 8C). Minor and major: anterior margin of clypeus of minor worker rounded and triangular, with anterolateral corner right-angled for C. maintikibo. Pronotum sometimes narrowly rounded or shouldered (C. ellioti, C. maintikibo) (Fig. 8B). In profile, dorsal outline of mesosoma evenly convex (C. ellioti, C. maintikibo) or slightly concave at the level of metanotal suture (C. andrianjaka) (Fig. 8D) or with impressed metanotal suture (C. maintilany). Humeral angles of pronotum well-marked. Petiole squamiform to nodiform. Integument finely imbricate to coarsely reticulate-foveolate. Pubescence long and distinct, few pairs of standing hairs on vertex.
Remarks. The Camponotus ellioti species group is characterized by the following morphological features: mesosoma reddish, gaster dark brown to black; anterior clypeal margin projecting into broadly convex lobe; frontal carina short and diverging posteriorly, median carina absent; clypeus with short, rounded lobe; mandible subtriangular, apical tooth long; eyes large, elliptical, and break the lateral margin of head; in minor, head and occipital margin rounded, lateral head sides parallel before eyes.
Camponotus ellioti has been considered a member of subgenus Myrmopiromis mainly because of the dense pilosity on its gastral dorsum and the humeral angle, but after morphological investigation, we find that the color of the integument can be a good indicator for separating Malagasy Camponotus species.
Minor worker: in full-face view, head elongate, lateral borders straight, feebly diverging posteriorly; posterior margin more or less convex (Fig. 9A).
Major worker: in full-face view, head rectangular, posterior and lateral borders slightly convex (Fig. 9D).
Minor and major worker: with the head in full-face view, anterior clypeal margin with an angular lobe, triangular or rectangular with rounded anterolateral angle; clypeus with median carina. In profile, dorsal line of mesosoma forms a continuous line, anterolateral corner of pronotum rounded. Petiolar node compressed anteroposteriorly and tapering dorsally; anterior face generally convex and posterior face straight. Posterior portion of the head, mesosomal dorsum, and the lateral margin of the propodeal declivity with scattered, whitish, erect hairs (Fig. 9B,D).
Remarks. The Camponotus madagascarensis species group possesses the same characters as the Camponotus niveosetosus group, described in Rakotonirina et al. (2017); the name has been changed because these species are endemic to Madagascar. This group is morphologically well defined by the distinctive disposition of hairs on the gastral segment. Four species of this group, with the exception of C. ivadia, are historically recognized as members of subgenus Myrmopiromis. In this study, we conclude that C. madagascarensis are endemic to the Malagasy region, while C. niveosetosus, the most similar species, is from South Africa and possesses the following suite of different morphological characters: occipital margin almost straight and not distinctly convex so that the anterior margin of pronotum is distinctly roof-like in dorsal view, cuticle of the dorsum of head of major worker with distinctive sculpture pattern, mandible of major worker with seven teeth.
Minor worker: in full-face view, head oval, with broadly convex posterior margin and lateral sides almost straight (Fig. 10A).
Major worker: in full-face view, head rectangular, posterior and lateral margins weakly convex (Fig. 10C).
Minor and major: with head in full-face view, clypeus carinate medially, its anterior margin forming a triangular lobe except for C. repens, which has a truncate anteromedian margin; dorsum of body covered with numerous slender, whitish, erect hairs and abundant, elongated pubescence. In lateral view, pronotum weakly convex with anterolateral portion marginate; promesonotal suture slightly depressed (Fig. 10B,D). Petiole nodiform except C. claveri. Head, mesosoma, and gaster black; legs generally much lighter in color than body. Integument shiny black. Remarks. Workers of the C. repens group can be diagnosed by the integument being black and shiny in appearance, and the presence of erect white setae along the entire dorsum, all of which are about the same length and transversely arranged, except in C. jjacquia, which has filiform and relatively medium-sized setae. Median portion of clypeus with longitudinal carina; dorsum of mesosoma covered with numerous, slender, erect hairs of about the same length; in lateral view, petiolar node higher than long. Metanotal spiracle obvious, surrounded by shallow depression, not as in C. christi species group. The best method to discriminate between the Camponotus repens species group and all other species stated in the Rakotonirina et al. (2018) Camponotus christi species group, is to compare the anterodorsum of pronotal margin and the mesosomal pilosity. In the C. repens group, in dorsal view, the pronotal disc is rectangular and pilosity consists of thick, whitish, erect hairs. By contrast, in the C. christi group, in dorsal view, the pronotal disc is semicircular, and pilosity consists of slender, filiform hairs.
Minor worker: in full-face view, head longer than broad, lateral margins nearly straight and slightly diverging posteriorly, posterior margin broadly convex (Fig. 11A). Major worker: in full-face view, head broader than long, lateral margin slightly convex (Fig. 11C).
Minor and major worker: clypeus not produced but with a rounded anterior margin. In profile, dorsal outline of mesosoma roughly straight and broken by metanotal suture (Fig. 11B,D); mesonotum and propodeum dorsum mostly rectangular and flattened in profile view. Ant black with opaque integument, pilosity suberect and white, pubescence short but distinct.
Remarks. The Camponotus robustus species group can be distinguished by the combination of the following characters: in profile, posterodorsal face of petiole flat; anterodorsal margin of pronotum not arcuate, as in C. edmondi species group, but with broadly rounded to tuberculate humeral angle. In dorsal view, propodeum dorsum rectangular with distinct lateral margin; it forms an obtuse angle with the declivitous face. Clypeus with short anterior rounded lobe, sometimes notched medially (C. ethicus). Mesosomal dorsum complex, metanotal suture impressed. This group includes three species, two of which have been previously described as part of the C. edmondi species group. However, from Rakotonirina et al. (2016), this group can be recognized with the combination of the following characters: dorsolateral margin of propodeum marginate or extending into a sharp ridge, propodeal declivity usually concave, ante- rolateral corner of pronotum most often marginate, forecoxa larger than the width of mesopleuron, and usually propodeal dorsum abruptly sloping down to the insertion of the petiole. In addition, this species group has a prominent pronotal suture which is large and arcuate, making the characteristic form of mesonotum oval in dorsal view; propodeum dorsum mostly reduced in length and inclined towards petiole; and petiolar face distinctly convex, with at least one face, in dorsal view, resembling a handheld fan. C. ethicus and C. robustus are now placed with Mayria and form a monophyletic group with Camponotus zoro based on the morphological traits cited above.
The NC Clustering dendrogram revealed 14 clusters (Fig. 12). Morphometric difference between Camponotus claveri and C. maintilany, C. darwinii and C. ursus are not detected by the clustering methods. Moreover, the distribution of each species, along with some unmeasurable features such as density of hair, integument opacity, and color of hair/integument/appendages mentioned at the species description, allowed us to consider them as separate species. Identification key to species for minor worker for Malagasy Camponotus (Mayria) Some couplets related to the 22 species that have been transferred into the subgenus Mayria are taken from Rakotonirina et al. (2016Rakotonirina et al. ( , 2017Rakotonirina et al. ( , 2018)).
1
With the combination of the following characters: head broadened posteriorly, head width equal to or slightly longer than pronotal width; occipital margin straight to evenly convex; in dorsal view, pronotum subrectangular with subparallel margins; mesosoma short and high, forming a rounded V in dorsal view; in profile, propodeum dorsum flat to slightly convex, never concave; petiolar node ranging from nodiform to squamiform; body integument opaque and dull with noticeable sculpture; pilosity consists of thick setae (Fig. 2 18
In profile, mesonotal dorsum strongly sloping down to the level of propodeum, maximum length of mesonotum about as long as distance between metanotal groove and propodeal spiracle (Fig. 29A); in dorsal view, lateral margin of mesonotum not well defined and converging gradually towards metanotal groove (Fig. 29B); head and mesosoma brown . Head, mesosoma, and petiolar node yellowish orange, gastral segments yellowish anteriorly and darkened posteriorly, very sparsely and finely pubescent; humeral angle rounded; petiole nodiform, node summit convex (Fig. 38A) ...maintikibo -Head, mesosoma, and petiolar node reddish to dark brown, gastral segments all black with dense, appressed, golden pubescence; humeral angle tuberculate; petiole squamiform with sharp edge (Fig. 38B) (CS: 1.85±0. Description of minor worker. Large-sized species. Absolute cephalic size (CS: 2.14±0.38, 2.00-2.33). In full-face view, head elongate (CWb/CL: 0.34±0.03; 0.32-0.35); posterior margin of head straight, lateral margin of head even straighter, feebly converging toward base of mandible. Eyes elliptical, sublateral, and placed next to the vertex (PoOC/CL: 0.08±0.02; 0.07-0.09). Mandibles triangular with six teeth. Clypeus strongly carinate throughout its length, with produced, angular, anterior margin, lateral corner angulate and median portion a rounded triangle (ClyL/GPD: 0.25±0.05, 0.22-0.28). Antennal scape longitudinally flattened, its base scarcely enlarged (SL/CS: 0.44±0.04, 0.42-0.45). In lateral view, mesosoma short and high, its dorsum smooth- on posteroventral head surface, scarcely distributed on mesosoma dorsum but much closer on declivity; decumbent, spatulate, whitish hair on the ventral section of gena, below the antennal socket, on the posterolateral margin of clypeus, promesonotum dorsum, petiolar node, and gastral tergite; suberect and decumbent whitish hairs present on gastral segment. Entire body blackish except the mandible testaceous to reddish.
Description of major worker. With characteristics of minor workers, except: head more cordate (CS: 3.32±0.21, 2.94-3.55) (CWb/CL: 0.96±0.01, 0.95-0.99); lateral margins slightly convex and tapering anteriorly. Eyes elliptical, smaller compared to head size (EL/CS: 0.20±0.01; 0.20-0.21), placed dorsally next to posterior margin of frontal carina. Antennal scape short and not exceeding posterior margin of head (SL/CS: 0.81±0.04; 0.76, 0.88); mandible more robust. Mesosoma and pilosity same as minor worker.
Distribution and biology. This new species is only known from Antsaraingy, a littoral forest, from 66 to 90 m in elevation, located in the northern portion of Madagascar (Fig. 70A). Nests were found underground and inside termite mounds; four colonies were collected to represent this species.
Discussion. Despite its large size, Camponotus antsaraingy may be confused with Camponotus mita and Camponotus voeltzkowii because in these three species the propodeal face of the propodeum is more reduced than the declivitous face. In C. antsaraingy the antennal segments are entirely flattened while in C. mita only the basal half is flattened; C. voeltzkowii has circular antennal segments. The pubescence on the gastral segment consists of thick, decumbent hair of the same type as the pilosity in C. antsaraingy, with short and filiform hairs on the other two species.
Etymology. The species epithet is in reference to the type locality. two major workers, same data as lectotype, but specimen coded respectively as CASENT0101195 and CASENT0101380 (MHNG). Camponotus (Myrmopiromis) darwinii r. rubropilosus var. robustior Wheeler, 1922Wheeler, : 1051;;Emery, 1925: 128;Bolton, 1995: 121. Syn. nov.
Body integument matte black, finely reticulate-punctate; mesosoma and/or gastral dorsum with dense, long, and yellowish pilosity and pubescence; anterior clypeal margin with short, rounded, rectangular lobe; petiole squamiform. Description of minor worker. Medium size. Absolute cephalic size (CS: 1.45±0.28; 1.22-1.63). In full-face view, head more quadrate (CWb/CL: 0.37±0.03, 0.35-0.41); posterior margin of head almost flat, lateral margin of head slightly convex and tapering to front. Eyes circular, placed midway from the occipital corner (PoOC/ CL: 0.08±0.02; 0.06-0.09). Mandibles triangular with six teeth. With head in fullface view, anterior clypeal margin with a short, rounded, rectangular lobe; less convex in lateral view, not medially marginate (ClyL/GPD: 0.25±0.04, 0.23-0.28). Antennal scape circular and long, surpassing the occiput by the length of two basal funiculi (SL/ CS: 0.40±0.04; 0.37-0.43). In lateral view, anterodorsal corner of pronotum rounded, mesosomal dorsal outline feebly and rather evenly arcuate above, dorsal face of propodeum generally longer than declivitous face (MW/ML: 0.21±0.02; 0.19-0.23; MPD/ML: 0.24±0.03; 0.20-0.26). In lateral view, petiolar node cuneate, anterodosum straight basally and inclined dorsally to reach the flattened posterodorsal margin. Tibia tubular, not prismatic, its extensor profile with suberect setae. Head dorsum is matte, finely reticulate-punctate with sparse punctures on malar area; mesosoma striate in lateral view; in dorsal view, petiole, mesosoma, and appendages less shiny, finely reticulate-striolate except the abdominal sternite, which is more matte. Mandible finely reticulate-punctate with sparse oblique punctures. Hairs yellowish white, orange, or tawny colored, erect hairs usually abundant, at least on gastral tergite, often on vertex, occipital portion, and thorax dorsum; petiolar node with 4 pairs of hairs on its dorsum. Shiny black, with mandible and basitarsus reddish to brownish.
Description of major worker. Characteristics of minor workers, except: head much wider posteriorly (CS: 2.14±0.26, 1.70-2.47; CWb/CL: 1.04±0.06, 0.94-1.20); lateral margins convex and converging towards base of mandible. Eyes semicircular, placed dorsolaterally midway from the occipital margin (PoOC/CL: 0.22±0.01, 0.19-0.24). Clypeus much more quadrate in full-face view (ClyL/GPD: 0.72± 0.09, 0.48-0.81), its anterior margin slightly concave medially. Antennal scape short, just surpassing the occipital border (SL/CS: 0.83±0.08, 0.75-0.99). Dorsal outline of mesosoma forms a continuous line so that the basal portion of propodeum forms an obtuse angle with the declivity. Lateral portion of head finely reticulate above, punctate to finely areolate below.
Distribution and biology. Camponotus darwinii is a famous species which is distributed widely across central of Madagascar, with a range extending from Ankazobe to Mount Papango, PN Befotaka Midongy (Fig. 58A). It has been collected from a wide variety of different habitats, including rainforest, montane rainforest, Uapaca forest, and Uapaca woodland as well as disturbed habitats such as urban and garden areas and cultivated land (Tavy). The species has also been found nesting inside Melia azedarach and Phellolophium madagascariense. Habitat elevations ranged from 1069-2150 m.
Discussion. Camponotus darwinii should be separable from most similar species in this group by the combination of the following characters: presence of dense hairs on mesosoma and/or gastral dorsum, mandible and malar area distinctly reddish in color, propodeal and declivitous faces unequal in length. This species is much closer to C. ursus but differs in the density of mesosomal hairs and the aspect of the petiole. In addition, C. darwinii is much more robust than C. ursus.
Additional material examined. Province Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815 Paratypes. Three workers with the same data as holotype: collection code: BLF35657, specimen code: CASENT0724324; collection code: BLF35614, specimen code: CASENT0724356; collection code: BLF35615, specimen code: CASENT0724357 (CAS).
Worker diagnosis. Integument black; head, mesosoma, and gastral dorsum clothed with dense, whitish hairs; anterior margin of clypeus with short rectangular lobe; dorsal outline of mesosoma almost flat; dorsum and declivitous faces of propodeum unequal in length and form an obtuse angle.
Description minor worker. Medium-sized species. Absolute cephalic size (CS: 1.45±0.24, 1.31-1.66). In full-face view, head more quadrate (CWb/CL: 0.36±0.02; 0.35-0.38), posterior margin of head and lateral margin weakly convex. Eyes circular, placed laterally close to lateral head margin (PoOC/CL: 0.08±0.01; 0.07-0.09). Mandibles triangular with six teeth. Clypeus convex, its anterior portion projected into a rectangular lobe. Antennal scape circular and long (SL/CS: 0.40±0.03; 0.37-0.42). In lateral view, mesosoma low and long, its dorsum smoothly arcuate, metanotal suture obsolete. Humeral angle rounded. Propodeal dorsum about the same length as declivitous face (MPH/ML: 0.18±0.02; 0.18-0.20). In lateral view, petiole squamiform with blunt edge, anterior face short and converging to the highest point of the petiole, posterior face straight. Ventral face of femora marginate laterally, the outer margin of tibia with long, whitish setae. Head and mesosoma coarsely reticulate-punctate, legs and gastral tergite finely imbricate. Mandible finely foveolate with sparse punctures. Pilosity and pubescence whitish, the former sparse and suberect, especially on occipital portion, frontal area, and mesosoma, though most conspicuous on the thoracic dorsum. On the gaster, the suberect hairs are long and abundant, overlapping with appressed hairs, pubescence inconspicuous and short. Petiolar node fringed with one row of whitish hair from the level of petiolar spiracle. Head including scape, mesosoma, and gaster dark brown to black, legs brownish, less dark than mesosoma.
Description of major worker. Characteristics of minor workers, except: head much wider posteriorly (CS: 1.98±0.30, 1.78-2.32; CWb/CL: 0.99±0.01, 0.98-1.00); lateral margins convex and converging towards base of mandible. Eyes semicircular, placed dorsolaterally midway from the occipital margin (PoOC/CL: 0.23± 0.01, 0.22-0.23). Clypeus much more quadrate in full-face view (ClyL/GPD: 0.72±0.02, 0.69-0.73), its anterior margin slightly concave medially. Antennal scape short, just surpassing the occipital border (SL/CS: 0.90±0.14, 0.75-1.00). Dorsal outline of mesosoma forms a continuous line so that the basal portion of propodeum forms an obtuse angle with the declivity. Lateral portion of head finely reticulate above, punctate to finely areolate below. (R. Harin'Hala) (CAS). Province Mahajanga: Parc National de Namoroka, 16.9 km 317° NW Vilanandro,45.31,100 m,tropical -13.291667, 48.258335, 146 m (Forel), AntWeb CASENT0101425 (MNHN). Paralectotype minor worker, same data as lectotype but specimen coded as CASENT0101373 (MHNG). Camponotus (Myrmobrachys) nossibeensis Forel, 1912: 91;1914: 270. Camponotus (Myrmepomis) nossibeensis Emery, 1920: 258. Camponotus (Myrmopiromis) nossibeensis Wheeler, 1922Wheeler, : 1052;;Emery, 1925: 129;Bolton, 1995: 114, 131.
Worker diagnosis. Integument matte black; body a rust brown color, relatively dense on entire dorsum; anterior margin of clypeus with short rectangular lobe; dorsal outline of mesosoma weakly arcuate; declivitous face of propodeum longer than its dorsum.
Description of minor worker. Large-sized species. Absolute cephalic size (CS: 2.24 0.46; 1.92-2.70). In full-face view, head slightly longer than broad, narrower in front than behind, with nearly straight posterior and lateral margins. (CWb/CL: 0.37±0.03; 0.35-0.42). Eyes more elongate, elliptical, not breaking outlines of head (PoOC/CL: 0.08±0.02; 0.06-0.09). Mandibles triangular withsix teeth. Clypeus carinate, anterior clypeal margin entire, projecting into a rectangular lobe with an obtuse anterolateral angle (ClyL/GPD: 0.28±0.05, 0.24-0.32). Antennal scape long, surpassing the occiput by the length one funiculus (SL/CS: 0.39±0.05; 0.33-0.41). In lateral view, mesosoma short, robust, and depressed; its dorsal outline arcuate; promesonotal suture large, distinctly shiny, and glabrous; humeral angle distinctly rounded; metanotal suture obsolete; propodeal dorsum slightly convex in profile and distinctly shorter than the declivitous face into which it passes via a rounded obtuse angle (MW/ ML: 0.22±0.03; 0.20-0.24; MPH/ML: 0.21±0.05; 0.18-0.24). In lateral view, petiolar node squamiform, anterior face confused with its dorsal face, node summit truncate, rounded triangle in dorsal view. Head, mesosoma, and gastral tergite finely and densely punctate. Mandible finely and longitudinally striate with sparse punctuation. Hairs golden yellow, erect to suberect, abundant, bending forward at its midlength on occipital region and mesosomal dorsum, but more abundant on mesonotum and propodeum dorsum; sparse, stiff, pointed, and a different length on gastral tergites. Petiole along its border with a fringe of golden, yellowish hairs not the same length. Body entirely black and a little shiny; masticatory margin reddish to dark brown.
Description of major worker. With characteristics of minor worker, except: head much wider posteriorly (CS: 3.05±0.41, 2.35-3.67; CWb/CL: 1.02±0.04, 0.96-1.08), lateral margin almost straight and tapering anteriorly to the mandibular insertion. Eyes elliptical, moderately sized compared to the head (EL/CS: 0.20±0.01, 0.1--0.23), placed dorsally next to the vertex (PoOC/CL: 0.23± 0.02, 0.18-0.27). Clypeus more rectangular (ClyL/GPD: 0.84±0.05; 0.73-0.91). Antennal scape just surpassing the occipital margin (SL/CS: 0.77±0.08; 0.68-0.91). Mandible strongly built with five large teeth, apical tooth sharp and long, and remainder decreasing in size to the basal margin. Mesosoma, pilosity, and pubescence same as minor worker. Head shiny black; anterior portion of head, including frontal area, finely alveolate punctate.
Distribution and biology. Camponotus nossibeensis is an endemic species mostly known from the northern part of Madagascar, from Nosy Faly to Ambanja. This species seems very adaple because it has been collected from both humid and tropical dry forest, at altitudes 7-780 m (Fig. 58C). The data indicate that individual workers forage on the ground and on lower vegetation, while nests are mostly found in dead twigs above ground, rotten logs, dead tree stumps, and underground.
Discussion. The combination of large size, big eyes, vertical declivitous face of propodeum, dense pilosity on mesosoma and gaster, and alveolate sculpture make C. nossibeensis a very distinctive species.
Additional material examined. Province Antsiranana: Forêt Ambato, 26.6 km 33° Ambanja, -13.4645, 48.55167, 150 m, rainforest (B.L. Fisher) (CAS); Nosy faly, Tafiambotry,35.3 km N Ambanja,48.48775,7 m,littoral secondary vegetation (BLF) (CAS); Sahamalaza Peninsula, Forêt d' Anabohazo, 21.6 km 247° WSW Maromandia, -14.30889, 47.91433, 120 m, tropical Wheeler, 1922Wheeler, : 1052;;Emery, 1925: 129;Bolton, 1995: 119, 131. Camponotus radovae-darwinii Forel, 1891: 46. Lectotype minor worker, present designation, Madagascar, Province Antananarivo (Camboué), AntWeb CASENT0101367 (MHNG). Paralectotype one worker with the same data as lectotype but specimen coded as CASENT0101115 (MHNG). Camponotus (Myrmopiromis) radovae-darwinii Wheeler, 1922Wheeler, : 1053;;Emery, 1925: 129;Bolton, 1995: 119. Syn, nov.
Integument matte black with sparse, yellowish brown hairs arranged in a series of transverse rows, pubescence reduced to absent on mesosomal dorsum. Anterior clypeal margin with a distinct produced rectangular lobe. Petiole cuneate in profile. Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 1.59±0.22, 1.43-1.77). In full-face view, head longer than broad, narrower in front than in back, with feebly convex occipital and lateral sides (CWb/CL: 0.37±0.03; 0.35-0.40). Eyes circular, placed much closer to the lateral sides (PoOC/CL: 0.08±0.01, 0.07-0.09). Mandibles triangular with six teeth. With head in full-face view, clypeus not carinate, its anterior margin produced, forming a short rectangular lobe with a sharp, lateral angle. Antennal scape circular, long, surpassing the occiput by the length of one basal funiculus (SL/CS: 0.39±0.03, 0.37-0.41). In lateral view, anterodorsal corner of pronotum broadly rounded, mesosomal dorsal almost straight, propodeal dorsum shorter than declivity, both straight in profile and meeting at a blunt angle (MW/ML: 0.21±0.02, 0.20-0.23; MPD/ML: 0.24±0.02, 0.22-0.26). Promesonotal and metanotal sutures form bold, polished lines. In lateral view, petiolar node cuneate, with a short, vertical, anterior face which reaches toward the flattened posterior faces. Head and mesosoma finely alveolate, gaster more finely strigulate. Mandible finely foveate with sparse punctures. Pilosity of dorsal head and body consists of fine and pointed brownish yellow hairs, long and suberect on frontal area and mesosomal dorsum; mostly short and sparse on gastral tergites; petiolar dorsum edge with 4 to 5 pairs of brownish yellow hairs. Body entirely matte black, apical portion of mandible, basitarsi, and funiculus reddish brown.
Description of major worker. Characteristics of minor workers, except: head somewhat cordate, with nearly straight posterior margin and slightly convex lateral sides (CS: 2.76±0.12, 2.59-2.85; CWb/CL: 1.05±0.02, 1.02-1.07). Eyes elliptical, placed dorsally midway from the occipital margin (PoOC/CL: 0.24±0.01, 0.23-0.25). Clypeus much more quadrate in full-face view with truncate anterior margin (ClyL/GPD: 0.76±0.13, 0.57-0.86). Antennal scape short, just surpassing the occipital border (SL/CS: 0.74±0.03, 0.72-0.79). Dorsal outline of mesosoma interrupted by pronotal and mesonotal sutures, basal portion of propodeum forms an obtuse angle with the declivity. Lateral portion of head finely punctate, frontal area and vertex with deep punctures.
Distribution and biology. Camponotus radovae is mainly distributed in the southern part of Madagascar in Zombitse National Park (Fig. 58D). This species is encountered in native forest habitats such as deciduous dry forest, tropical dry forests, gallery forests, and spiny forests and thickets. It was sampled mostly from Malaise traps and by beating lower vegetation, on the ground, and twice on leaf litter at altitudes 20-840 m. Wheeler, 1922Wheeler, : 1051;;Emery, 1925: 128. Worker diagnosis. Integument dark brown with rust or red-brown suberect pilosity. Anterior clypeal margin with rectangular projection. Petiole scale thin with sharp border.
Description of minor worker. Medium-sized. Absolute cephalic size (CS: 1.82±0.37; 1.53-1.98). In full-face view, head slightly longer than broad, a little narrower in front than in back, with straight occipital and feebly convex lateral margins (CWb/CL: 0.36±0.03; 0.34-0.38). Eyes circular, placed much closer to the lateral sides (PoOC/CL: 0.07±0.01; 0.07-0.08). Mandibles triangular with sixteeth. With head in full-face view, anterior clypeal margin produced, entire, and angularly project-ing in the middle. Antennal scape circular, long, surpassing the occiput by the lengths of two basal funiculi (SL/CS: 0.41±0.05; 0.38-0.44). In lateral view, anterodorsal corner of pronotum broadly rounded, mesosomal dorsum moderately arcuate, propodeum dorsum a little shorter than the declivity, each straight in profile and meeting at an obtuse angle (MW/ML: 0.21±0.02, 0.20-0.22; MPD/ML: 0.24±0.03; 0.22-0.26). In lateral view, petiolar node wedge-shaped, with feebly convex anterior and flattened posterior faces. Ants entirely reticulate-punctate. Mandible finely striolate with sparse punctures. Pilosity and pubescence brownish yellow, the former long and curved forward especially on vertex, frontal area, and mesosomal dorsum; mostly short and sparse on gastral tergites; petiolar dorsum edge with four or five pairs of brownish yellow hairs. Body color matte black, apical portion of mandible reddish.
Description of major worker. Characteristics of minor workers, except: head somewhat cordate, with nearly straight posterior margin and slightly convex lateral sides (CS: 2.68±0.36, 2.26-3.15; CWb/CL: 1.05±0.04, 0.99-1.11). Eyes elliptical, placed dorsally midway from the occipital margin (PoOC/CL: 0.23±0.01, 0.22-0.25). Clypeus much more quadrate in full-face view with truncate anterior margin (ClyL/GPD: 0.82±0.07, 0.73-0.93). Antennal scape short, just surpassing the occipital border (SL/ CS: 0.75±0.03, 0.71-0.79). Dorsal outline of mesosoma interrupted by pronotal and mesonotal sutures, basal portion of propodeum forms an obtuse angle with the declivity. Lateral portion of head finely punctate, frontal area and vertex with deep punctures.
Distribution and biology. Camponotus themistocles has been collected at three localities in southern Madagascar (Fig. 58E): two are littoral forest and one is gallery forest. All sites are at low altitudes from 10 to 30 meters. Camponotus themistocles is a ground nester; most collections have been made from sifted litter, rotten logs, and dead twigs above the ground.
Discussion. Camponotus themistocles differs from C. radovae by its larger size and the sculpture of the lateral sides of its mesosoma, which is finely reticulate-punctate in the former and widely striolate in the later. In addition, in lateral view, the petiole scale of C. themistocles has an acute summit but is blunt in C. radovae. Pubescence is long and abundant in C. themistocles, and short and dilute in C. radovae. Morphometrical analysis reveals that C. themistocles and C. ellioti are the same size, but differ in integument coloration and pilosity pattern on the gastral tergite.
Additional material examined. Province Toliara: Forêt de Petriky, 12.5 km W 272° Tolagnaro, -25.06167, 46.87, 10 m, littoral rainforest (B.L. Fisher) (CAS); Forêt Mandena 8.5 km N Tolagnaro,47.0025, 20 m, littoral rainforest (BLF) (CAS); Mandena, 8.4 km NNE 30° Tolagnaro,47.00167, 20 m, littoral rainforest (B.L. Fisher) (CAS); Ranobe, -23.04085, 43.61012, 30 m, gallery forest (Frontier Wilderness Project) (CAS).
Camponotus ursus Forel, 1886: ci. Lectotype minor worker, present designation, Madagascar (Forel), AntWeb CASENT0101374 (MHNG) [examined]. Paralectotype, one minor worker same data as lectotype but specimen coded as CASENT0101375 (MHNG). Camponotus (Myrmobrachys) ursus Forel, 1912: 91;1914: 271. Camponotus (Myrmepomis) ursus Emery, 1920: 258. Camponotus (Myrmopiromis) ursus Wheeler, 1922Wheeler, : 1053;;Emery, 1925: 129;Bolton, 1995: 128, 131.
Worker diagnosis. Integument shiny black; anterior margin of clypeus with short, rectangular lobe; dorsum of mesosoma with dense, decumbent, golden-yellow setae; gastral dorsum with widely distributed, short, subdecumbent setae.
Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 1.54±0.19; 1.43-1.64). In full-face view, head somewhat longer than broad, narrower in front than behind, with slightly convex lateral and posterior sides (CWb/CL: 0.37±0.02; 0.36-0.38). Eyes elliptical, sublateral at midpoint from the lateral sides (PoOC/CL: 0.07±0.02; 0.06-0.08). Mandibles triangular with six teeth. Clypeus not carinate, produced into a short, rectangular lobe. Antennal scape long, surpassing the occiput by the length of one basal funiculus. (SL/CS: 0.38±0.02; 0.37-0.39). In lateral view, dorsal contour of mesosoma smoothly convex, humeral angle broadly rounded; in dorsal view, mesonotal suture distinct but not impressed, mesonotal suture obsolete so that mesonotum and propodeum are fused together, propodeum with convex base and sloping declivity (MPH/ML: 0.18±0.02; 0.16-0.19). Petiole narrow, cuneate in profile, with short anterior face tapering dorsally to the flattened posterior face, its border rather sharp, produced upwards as a blunt angle in the middle. Head dorsum finely reticulatepunctate with shallow, sparse punctures; occipital region transversally striate-reticulate; lateral face of mesosoma, declivitous face, and petiolar face finely strigulate; legs finely reticulate; dorsum of mesosoma finely reticulate-striate-punctate with sparse excavation from which one suberect setae arises. Dorsum of gastral tergite finely striate-reticulate transversally, with sparse, small punctures. Mandible finely rugose with sparse, large punctures. Hairs golden yellow, the former abundant, long, and bending forward on entire mesosoma dorsum; suberect, short, and sparsely distributed on gastral tergites; the latter short and conspicuous on the abdominal segment, five pairs of erect hairs present on vertex. Body shiny black; scape, two basal funiculi, mandible, tarsi, and tibiae reddish.
Description of major worker. Characteristics of minor workers, except: head as broad as long, with occipital and lateral margin almost straight (CS: 2.14±0.15, 2.00-2.39; CWb/CL: 0.89±0.07, 0.84-1.03). Eyes circular, placed dorsally next to the vertex (PoOC/CL: 0.23±0.01, 0.21-0.24). Anterior clypeal margin forms a short, rounded lobe (ClyL/GPD: 1.03±0.41, 0.73-1.78). Antennal scape short, just reaching the occipital border (SL/CS: 0.82±0.03, 0.78-0.86). Dorsal outline of mesosoma almost flat, propodeum dorsum naked and the same length as the sloping declivity.
Distribution and biology. Camponotus ursus is found in two different habitats: primary forest in the eastern portion and urban/garden areas in the central highlands of Madagascar (Fig. 58F). It is found foraging on low vegetation or inside branches above the ground. It occurs at altitudes above 1,200 meters.
Discussion. Camponotus ursus is recognizable within the C. darwinii species group on the basis of its distinct mesosomal pilosity. In addition, it is the only species with reddish brown basitarsi. However, C. ursus is unlikely be confused with C darwinii for several reasons. First, the body of the latter is much larger. Second, the head and gastral segment of C. ursus are covered with fine, short, sparse, and yellowish setae, giving the ant a glossy appearance that is completely different than that of C. darwinii.
Additional material examined. Province Antananarivo: Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (MHNG); Ankazobe, -18.31617, 47.11583, 1241 m (BLF) (CAS). Province Toamasina: Manakambahiny Atsinanana, -17.75, 48.71667 (A. Pauly) (CAS). Worker diagnosis. Camponotus chrislaini can be easily distinguished from the other members of the efitra group by its mostly smooth and shiny sculpture; head more or less globose in full-face view; sigmoid frontal carina, and wavy propodeum dorsum. Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 0.95 ± 0.17; 0.87-1.02). In full-face view, head ovoid (CWb/CL: 0.34±0.03; 0.32-0.35); posterior margin rounded, lateral margin of head convex. Eyes circular, protruding, placed midway from the occipital corner (PoOC/CL: 0.12±0.03; 0.11-0.13). Mandibles triangular with five teeth. Clypeus transversally trapezoidal, its anterior border broadly rounded, not angular. With head in full-face view, anterior clypeal margin rounded, not projecting (ClyL/GPD: 0.26±0.03, 0.25-0.29). Antennal scape flattened and lobulated at the base, long, surpassing the occiput by the length of three basal funiculi (SL/CS: 0.46±0.03; 0.45-0.48). In lateral view, anterodorsal corner of pronotum tuberculate, pronotal suture distinct, mesonotal suture distinctly impressed so that the anterior por- tion of propodeum is raised to a blunt edge in profile, and suture forms a concavity (MW/ML: 0.22±0.02, 0.21-0.23; MPH/ML: 0.17±0.02, 0.17-0.18). In lateral view, petiolar node nodiform, higher than long. Entire body, including mandible, smooth and shiny. One pair of standing hairs on angle formed by mesonotal dorsum and propodeum, head with scattered, subdecumbent, filiform brown hairs, scape with decumbent setae. Ant brownish; mandible, clypeus, and pronotum light brown; mid-and forecoxae, apical femora, basal tibiae, and apical portion of third and fourth tergite whitish.
Description of major worker. Characteristics of minor workers, except: head quadrate with straight margins (CS: 1.67±0.06, 1.57-1.74; CWb/CL: 0.92±0.03, 0.90-0.96). Eyes circular, closed to the lateral margin (PoOC/CL: 0.29±0.02, 0.27-0.31). In profile, clypeus distinctly truncate (ClyL/GPD: 0.92±0.03, 0.86-0.94), its anterior margin with a short concavity in full-face view. Antennal scape same as minor but short, just reaching the occipital border (SL/CS: 0.77±0.04, 0.74-0.83). Promesonotum forms a unique dome followed by an impressed mesonotal suture, dorsum of propodeum slightly concave near the propodeal angle. Anterior portion of head with widely spaced large punctures, a single puncture in the center of the vertex. Filiform brownish hair present on head and mesosoma dorsum, more abundant on malar area.
Distribution and biology. Camponotus chrislaini is restricted to the northern portion of Madagascar (Fig. 70B). This newly identified species prefers dry (tropical dry forest, dry forest, disturbed dry forest) as well as wet (montane forest and rainforest) habitats at moderate altitudes (below 1000 m). It was primarily collected from rotten logs, sifted litter, inside trees, or on branches above ground.
Discussion. The new species is easily distinguishable from the Mayria species, but morphologically is most similar to the unidentified species Camponotus MG098 from subgenus Myrmonesites. The main character used to distinguish them is the form of the petiolar scale, as in the former the petiole is much thinner and its dorsal face forms a straight surface, while in the latter the petiole is thickened with the posterior face concave in dorsal view. Camponotus MG098 is also covered with plentiful setae which are scarce and widely distributed in C. chrislaini. The presence of C. chrislaini and C. MG098 in six localities (Sakaramy, Binara, Antsahabe, Analamerana-Ankavanana, Galoko, and Mont Kalabenono) suggests that they are reproductively isolated.
Additional material examined. Province Antsiranana: Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical Betsiaka, -13.11833, 49.23, 425 m, rainforest (B.L. Fisher) (CAS); Res. Analamerana,49.37383,225 m,tropical Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (FGAT) (CAS); Reserve Spéciale de l'Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest (FGAT) (CAS); Ampasindava, Forêt d'Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (FGAT) (CAS).
A description of Camponotus maintikibo can be found in Rakotonirina et al. (2016) February 2010 (ARA), collection code: ARA0306, specimen code: CASENT0243690 (CAS). Paratype. one worker with same data as holotype but specimen coded as CASENT0243954 (CAS).
Worker diagnosis. Camponotus andrianjaka can be easily distinguished from other species of the ellioti group by the following character set: body bicolored, head and mesosoma reddish brown, gaster dark brown to black; anterior clypeal margin rounded, not produced; propodeum dorsum slightly concave at the level of metanotal suture; petiole nodiform and surmounted with standing, whitish hairs on its posterior margin.
Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 0.84±0.07; 0.81-0.88). In full-face view, head subovate, usually somewhat longer than broad, posterior cephalic margin roundly convex; lateral margin of head straight and tapering to front (CWb/CL: 0.31±0.01, 0.30-0.32). Eyes subelliptical, protruding, its border smoothly aligned to lateral margins of head (PoOC/CL: 0.08±0.01, 0.08-0.09). Mandibles triangular with six teeth. Clypeus carinate, anterior clypeal margin with rounded triangular projection (ClyL/GPD: 0.24±0.01, 0.23-0.25). Antennal scape long, surpassing the occiput by the length of two basal funiculi (SL/CS: 0.36±0.04, 0.35-0.38). In lateral view, mesosoma without anterolateral margination, promesonotum convex, mesopropodeal suture shallowly impressed, propodeum rather angulate with slightly concave basal and declivitous surface (MW/ML: 0.19±0.02, 0.18-0.21, MPH/ML: 0.15±0.01, 0.14-0.15). In lateral view, petiole nodiform, higher than long, its short anterior face rounding gradually to the convex posterior faces. Head, especially anterior portion, finely reticulate-areolate, gaster and mesosoma finely imbricate. Mandibles finely reticulate-punctate. Hairs whitish, one pair present on vertex, middle of mesonotum, and propodeum dorsum; three to four pairs on each side of lateral corner of propodeum; four pairs transversely arranged on petiolar posteromargin; ground pubescence delicate. Head and mesosoma reddish, legs light brown, basal face of first gastral segment light brown and the remainder of the segment dark brown to black.
Description of major worker. Characteristics of minor workers, except: head as long as broad, with rather straight posterior and lateral borders (CS: 1.15±0.05, 1.12-1.20, CWb/CL: 0.84±0.01, 0.84-0.85). Eyes circular, placed dorsally next to lateral borders (PoOC/CL: 0.26±0.03, 0.23-0.27). Anterior clypeal margin forms a rounded rectangular lobe (ClyL/GPD: 0.89±0.05, 0.83-0.92). Antennal scape short, just reaching the occipital border (SL/CS: 0.70±0.02, 0.69-0.72). Mesosoma the same as minor worker.
Distribution and biology. Camponotus andrianjaka is a very distinctive species that appears endemic to Bismarckia woodland, shrubland, Uapaca woodland, and savannah woodland in the southern part of Madagascar (Fig. 70D). It can be found also on eucalyptus plantations and grassland. Altitude: 1300-1987 m.
Discussion. At first sight, Camponotus andrianjaka may be confused with another new species, Camponotus MG132, subgenus Myrmonesites, but can be separated easily with mesonotum that is gradually rounded in dorsal view; petiole higher than long, its dorsolateral margin surmounted with white hairs; presence of ground pubescence; and gaster all black. In addition, C. MG132 has been collected from rainforest and montane rainforest and absent from woodland.
Etymology. This new species is named after the collector Andrianjaka Ravelomanana. Additional material examined. Province Fianarantsoa: Ampandravelo III Non Protected Area, 10.72 km NE Ranohira, -22.53944, 45.51497, 869 m, shrubland (ARA) (CAS); Ampangabe III Non Protected Area, 21.26 km W Itremo, -20.6125, 46.60883, 1412 m, savannah woodland (ARA) (CAS); Ampangabe VI Non Protected Area, 21.16
Camponotus ellioti Forel, 1891: 37, 73. Holotype minor worker, Madagascar, collection Henri de Saussure. AntWeb CASENT0101382 (MHNG) (examined). Camponotus (Myrmepomis) ellioti Forel, 1912: 92;Emery, 1920: 258. Camponotus (Myrmopiromis) ellioti Wheeler, 1922Wheeler, : 1052;;Emery, 1925: 128;Bolton, 1995: 97, 131. Camponotus ellioti var. relucens Santschi, 1911: 133. Lectotype two workers, present designation, Madagascar, Toliara Province, Region du Sud-Est Fort-Dauphin (Ch. Alluaudi), AntWeb CASENT0101196 (minor); CASENT0101197 (major) (NHMB). Camponotus (Myrmopiromis) ellioti var. relucens Wheeler, 1922Wheeler, : 1052;;Emery, 1925: 129;Bolton, 1995: 120. Syn. nov. Diagnosis. Head and mesosoma reddish brown to dark brown, gaster always dark colored; anterior clypeal margin of clypeus with short, rectangular lobe; pronotal humeri tuberculate, dorsal outline of mesosoma arcuate; gastral tergite with short, decumbent, ochreous setae. Description of minor worker. Large-sized species. Absolute cephalic size (CS: 1.85±0. 55, 1.45-2.42). In full-face view, head trapezoidal, wider than length of pronotum in dorsal view; occipital margin slightly convex medially, and straight subparallel lateral margin of head (CWb/CL: 0.36±0.03, 0.33-0.39). Eyes elliptical, positioned at the midline of head (PoOC/CL: 0.08±0.01, 0.07-0.09). Mandibles short and triangular with six teeth. Clypeus trapeziform, less convex in profile, sharply carinate along its entire length, with a short, rectangular projection feebly sinuate in the middle (ClyL/GPD: 0.12±0.01, 0.11-0.13). Frontal carina less sinuate and divergent. Antennal scape long and surpassing the occipital corner (SL/CS:0.41±0.07;0.34-0.46). In lateral view, dorsal outline of mesosoma forms a smooth curve, anterodorsal corner of pronotum distinctly shouldered, propodeum rounded, its surface convex and not marginate laterally, declivitous face triangular in posterior view (MW/ML: 0.23±0.03, 0.21-0.25; MPH/ML: 0.18±0.03, 0.16-0.20). Petiolar node squamiform in profile, wider than long, anterodorsal face distinctly flat and inclined forward, posterodorsal face flat, node summit arcuate. First abdominal tergite with vertical anterior face. Entire body (frontal area, dorsal face of mesosoma, scape, and legs) entirely, finely, and densely reticulate-punctate and matte; lateral face of petiole and abdomen transversally striolate. Mandible finely reticulate-punctate and subopaque, with superimposed, abundant punctuation arranged transversally. Ground pilosity yellowish, short, sparse, and dilute on tibia and scape. Short white to yellowish standing hair present on entire dorsum, sparsely distributed on occipital region, regularly present on petiolar dorsum and onto declivity, absent on scape and tibia. Dorsum of gastral segment with appressed ochreous setae and whitish, suberect hairs arranged serially parallel and longitudinally. Head and mesosoma dark brown to reddish brown; mandible, basal half of scape, basitarsus, and gastral segment reddish.
Description of major worker. Characteristics of minor workers, except: head as long as broad, with rather straight posterior and lateral borders (CS: 4.13±0.16, 3.90-4.33; CWb/CL: 1.04±0.02, 1.01-1.07). Eyes circular, placed dorsally next to lateral borders (PoOC/CL: 0.28±0.01, 0.26-0.29). Anterior clypeal margin forms a rectangular lobe (ClyL/GPD: 0.87±0.05, 0.79-0.92). Antennal scape short, not reaching the occipital border (SL/CS: 0.61±0.02, 0.58-0.65). Promesonotal suture strongly marked.
Distribution and biology. Camponotus ellioti occurs in the dry forest regions of the southeast of Madagascar in habitats such as bush, coastal scrub, and coastal spiny bush on sandy soil, Euphorbia forest, spiny bush and thicket, gallery forest, and riparian scrub (Fig. 70C). It has been collected from sifted litter and Malaise traps. The altitudes of these localities range from 5 to 230 meters.
Discussion. Camponotus ellioti is a distinctive species recognizable by the submargined anterolateral corner of the pronotum and the gaster covering of thick, blunt hairs the color of rust. In the traditional classification, C. ellioti is considered a member of the C. darwinii species group based on the density and color of the gastral pilosity and the sculpture pattern; however, we propose color as a character difference between these two groups. same data as holotype but collection code: BLF26364, specimen code: CASENT0245172; collection code: BLF26385, specimen code: CASENT0245187 (CAS).
Worker diagnosis. Ants bicolored: head and mesosoma reddish brown, gaster dark brown to black; pronotum and mesonotum form a unique dome followed by an impressed metanotal suture, propodeum, and petiole angle with standing, whitish, spatulate hairs, head and gastral dorsum with sparse, fine, brown hairs; petiole squamiform.
Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 0.84±0.05, 0.81-0.86). In full-face view, head distinctly longer than broad, with con- vex occipital margin and subparallel lateral sides (CWb/CL: 0.33±0.02, 0.32-0.34). Eyes elliptical and break the lateral outlines of head (PoOC/CL: 0.09±0.01, 0.08-0.10). Mandibles triangular with six short, blunt teeth. Clypeus convex, its anterior border produced into rounded lobe (ClyL/GPD: 0.24±0.01, 0.22-0.25). Antennal scape long, surpassing the occipital margin (SL/CS: 0.40±0.03, 0.39-0.43). In lateral view, mesosoma without anterolateral margination, promesonotum distinctly convex, mesopropodeal suture deeply impressed so that propodeum dorsum is raised to the level of mesopropodeal suture, propodeum angular in profile, with unequal basal and declivitous faces, the former horizontal, the latter straight to evenly concave (MW/ ML: 0.20±0.01, 0.19-0.20; MPH/ML: 0.14±0.03, 0.12-0.15). In lateral view, petiole cuneate, its anterior face convex, its posterior face more flattened with entire border broad. Body finely imbricate. Hairs grayish, present only on vertex, one pair next to the pronotal suture, one pair on the middle of mesonotum dorsum, sparse on gastral tergite; enlarged, whitish hairs present on propodeal corner (two or three pairs) and on posteromargin of petiole; pubescence whitish and more dilute on gaster tergites. Head, mesosoma, and appendages reddish to brownish yellow; first gastral segment lighter colored than the remaining dark brown segments.
Description of major worker. Characteristics of minor workers, except: head apparently rectangular, with straight to feebly convex posterior margin and parallel lateral sides. (CS: 1.13±0.05, 1.09-1.18; CWb/CL: 0.90±0.02, 0.88-0.91). Eyes circular, placed dorsally next to the lateral margin (PoOC/CL: 0.26±0.01, 0.25-0.27). Anterior clypeal margin forms a short, rounded lobe (ClyL/GPD: 0.83± 0.00, 0.83-0.83). Antennal scape short, just reaching the occipital border (SL/CS: 0.77±0.03, 0.74-0.79). Dorsal outline of mesosoma not very convex and interrupted but pronotal and mesopropodeal suture present, the latter weakly impressed.
Distribution and biology. Camponotus maintilany occurs from 1300-1987 m in elevation and inhabits a montane environment in the central highlands, where it prefers mostly Uapaca forest or woodland, but is also found in savanna grassland and eucalyptus plantations (Fig. 70E). This new species has been collected by digging in soil, sifting leaf litter, and baiting with tuna fish or sardines. In addition, a few workers were found in Malaise traps.
Discussion. Camponotus maintilany is most readily distinguished from other members of the ellioti species group by a deeper metanotal groove and shinier sculpture. It may be confused with C. andrianjaka but can be separated by its impressed metanotal suture and truncate petiolar node.
Etymology. This species is derived from the coloration of the mesosoma and gaster. Worker diagnosis. Integument dark brown, apparently shiny; hairs on mesosoma and petiole blunt-tipped; fine, pointed hairs on head and gastral dorsum; anterior clypeal margin with straight rectangular lobe; dorsal outline of mesosoma nearly evenly arcuate; petiole higher than long.
Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 1.21±0.19, 1.13-1.30). In full-face view, head almost quadrate, with feebly convex posterior and lateral border (CWb/CL: 0.35±0.03, 0.33-0.36). Eyes elliptical, placed next to the lateral margins of head (PoOC/CL: 0.10±0.01, 0.10-0.11).
Mandibles triangular with six teeth. Anterior clypeal margin angularly produced, with straight lateral border and rounded anterolateral corner. Antennal scape long, surpassing the occipital corner by the length of two basal funiculi (SL/CS: 0.39±0.02, 0.38-0.40). In lateral view, pronotum with short anterolateral margination, mesosoma moderately convex, propodeum angular in profile, with unequal basal and declivitous faces, each straight in profile and meeting at a rounded obtuse angle (MW/ML: 0.21±0.03, 0.19-0.22; MPH/ML: 0.16±0.01, 0.16-0.17). In lateral view, petiole rather narrow, its anterior face convex and its posterior face flattened. Body finely imbricate. Hairs whitish; enlarged, whitish hairs with blunt tip on mesonotum, propodeum, and petiolar dorsum; filiform, pointed, grayish hair on head, pronotum, and gastral segment; pubescence white, short, and more abundant across entire dorsum. Head, mesosoma and gaster dark brown, femora, tibia, and funiculi brown, scape and basitarsi slightly lighter.
Description of major worker. Characteristics of minor workers, except: head as long as wide, posterior borders medially concave and subparallel, lateral sides diverging apically (CS: 1.93; CWb/CL: 0.93). Eyes circular, placed dorsally in the middle of the lateral borders (PoOC/CL: 0.28). Anterior clypeal margin forms a short, rounded, rectangular lobe, slightly concave medially (ClyL/GPD: 0.84). Antennal scape short, just reaching the occipital border (SL/CS: 0.69). Sculpture, pilosity, and color as in minor worker.
Distribution and biology. Camponotus ivadia has thus far been collected only in the Forêt d' Ampondrabe within Ankarana Reserve, a region with tropical dry forest located in the northern portion of Madagascar (Fig. 70F). All specimens were collected by beating vegetation.
Discussion. This species is easily recognized within the madagascarensis group as one of few back and shiny species with white standing pubescence.
Etymology. Worker diagnosis. Integument black; entire body with plentiful, erect setae; anterior clypeal margin produced into a rounded triangular lobe; dorsal outline of mesosoma slightly impressed at the level of metanotum; petiole long and low.
Description of minor worker. Medium-sized. Absolute cephalic size (CS: 0.99±0.14; 0.91-1.09). In full-face view, head almost quadrate (CWb/CL: 0.32 ± 0.02; 0.31-0.34); posterior margin evenly straight, lateral margin of head ante-rior to eyes slightly convex. Eyes circular, placed above midline of head (PoOC/CL: 0.10±0.02; 0.09-0.12). Mandibles triangular with six teeth. Clypeus carinate, its anterior border angularly produced in the middle (ClyL/GPD: 0.20±0.06, 0.16-0.23). Antennal scape long, surpassing the occiput by the length of one basal funiculus (SL/ CS: 0.34±0.02; 0.32-0.34). In lateral view, anterodorsal corner of pronotum rounded, mesonotum dorsum inclined toward nearly straight propodeum face (MW/ML: 0.19±0.02, 0.18-0.20; MPH/ML: 0.13±0.02, 0.12-0.14). In lateral view, petiolar node longer than wide, with flat summit, its anterior face leaning forward. Ants finely foveolate throughout, apparently shiny. Entire body, including mandibles and appendages, covered with plentiful, erect, whitish setae, uneven in length, some long. Head, mesosoma, and gaster dark brown to black; antennae, mandibles, tibiae, and tarsi reddish brown.
Description of major worker. Characteristics of minor workers, except: head much wider posteriorly (CS: 1.68±0.07, 1.61-1.76; CWb/CL: 0.91±0.02, 0.89-0.92); lateral margins convex and converging towards base of mandible. Eyes circular placed dorsolaterally midway from the occipital margin (PoOC/CL: 0.30±0.00, 0.29-0.30). Clypeus with a short rectangular lobe with rounded lateral angle (ClyL/ GPD: 0.85±0.03, 0.81-0.88). Antennal scape short, not reaching the occipital margin (SL/CS: 0.57±0.02, 0.56-0.59). Dorsal outline of mesosoma interrupted by sutures, metanotal suture forms a canal. Head dorsum and mesosoma coarsely reticulate-punctate, petiole finely striolate, gaster imbricate and shiny.
Distribution and biology. The known elevational range of Camponotus jjacquia extends from 43 to 923 m. Specimens have been collected on tree trunks, on low vegetation, in leaf litter, and under stones in Bismarckia woodland, deciduous forest, gallery forest, savannah woodland, spiny forest, and tropical dry forest (Fig. 72C). This species nests in the soil.
Discussion. The unique petiolar shape, the density of its pilosity, and the shape of its metanotal dorsum make C. jjacquia a very distinctive species.
Etymology. The new species is dedicated to Jean Jacques Rafanomezantsoa, who has been a Malagasy ant collector for more than twenty years.
Additional material examined. Province Fianarantsoa: Ampotoampoto III National Park, 7.91 km NW Ilakaka, -22.62944, 45.189, 919 m, savannah woodland (ARA) (CAS); Isalo IV National Park, 12 km SW Ranohira, -22.61472, 45.31304, 867 m, Bismarckia woodland (ARA) (CAS); Ampotoampoto IV National Park, 7.83 km NW Ilakaka, -22.62944, 45.1912, 923 m, savanna 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro,46.6455,300 m,tropical dry forest (FGAT) (CAS); Parc National d'Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550m, montane rainforest, Malaise trap, 22-26 January 2003 (BLF), collection code: BLF07152, specimen code: CASENT0490618 (CAS). Paratypes. Three workers: one worker with same data as holotype but specimen collected from Malaise trap and with collection code: BLF07155, specimen code: CASENT0049849; two workers from Madagascar, Province Fianarantsoa, Mampiarika IV Non Protected Area, 27.98 km SW Ambositra, -20.73528, 47.08382, 1486 m, Uapaca woodland, 03-07 February 2010 (ARA), collection code: ARA0492, specimen code: CASENT0168788 and CASENT0168787 (CAS).
Worker diagnosis. Integument shiny black; anterior clypeal margin with short, subtriangular lobe; erect hairs arranged in a transverse row on gastral segment, sparsely arranged on mesosoma dorsum; dorsum of mesosoma with promesonotal and mesometanotal sutures marked but not impressed; petiole squamiform.
Description of minor worker. Medium-sized species. Absolute cephalic size (CS: 0.89±0.08, 0.84-0.93). In full-face view, head as long as wide, posterior margin rounded, lateral margin straight and subparallel (CWb/CL: 0.33±0.02, 0.32-0.34). Eye circular, protruding, its border smoothly aligned to lateral margins of head (PoOC/CL: 0.10±0.02, 0.08-0.10). Mandibles triangular with six teeth. Clypeus carinate, its anterior margin produced medially into a subtriangular lobe (ClyL/ GPD: 0.23±0.07, 0.20-0.26). Antennal scape long, surpassing the occipital corner by the length of two-and-a-half basal funiculi (SL/CS: 0.39±0.03, 0.37-0.40). In lateral view, mesosoma without anterolateral margination, promesonotum not very convex, mesopropodeal suture impressed, propodeum angular with the base fully twice as long as the declivity, which is straight (MW/ML: 0.18±0.01, 0.18-0.19; MPH/ ML: 0.14±0.01, 0.14-0.15). Petiole cuneate in profile, its anterior face convex, and its posterior face flattened along entire broad border. Body finely imbricate. Hairs whitish, one pair present on vertex, middle of mesonotum, and propodeum dorsum; two pairs on each side of lateral corner of propodeum; three pairs on lateral margins on petiole. Head and mesosoma reddish, legs light brown, basal face of first gastral segment light brown and the remaining segment dark brown to black.
Description of major worker. Characteristics of minor workers, except: head as long as broad, with rather convex posterior borders and subparallel lateral sides (CS: 1.18±0.09, 1.12-1.24, CWb/CL: 0.86±0.01, 0.85-0.87). Eyes circular, placed dorsally next to lateral borders (PoOC/CL: 0.26±0.00, 0.25-0.26). Anterior clypeal margin forms a rounded rectangular lobe (ClyL/GPD: 0.88±0.05, 0.85-0.91). Antennal scape short, just reaching the occipital border (SL/CS: 0.75±0.04, 0.73-0.78). Mesosoma with the same form as minor worker. Sculpture on head coarsely reticulatepunctate anteriorly and tends to be striolate posteriorly.
Distribution and biology. Camponotus claveri occupies two different habitats: montane rainforest and open habitats such as Uapaca woodland, eucalyptus plantation, shrubland, and deciduous dry forest, plus urban areas (Fig. 72A). Their nests are found in soil, under stones, and on the ground. It has been collected at elevations of 830-1987 m.
Discussion. Camponotus claveri is similar to other species of the repens group, but differs in having a thinner petiolar node with sharp dorsal edge, and a rounded triangular anterior clypeal margin. Morphometric analysis combined C. claveri and C. maintilany in the same cluster, meaning that they are the same size but have different qualitative morphological traits. Their distributions are quite distant but sympatric at Alasora, situated in the central highlands.
Etymology. This species is named after the collector Claver Marotafika Randrianandrasana.
Additional material examined. Province Antananarivo: Alasora, -18.96245, 47.58925, 1434 m, eucalyptus plantation (BLF) (CAS); Antaponimanadala III Non Protected Area, 6.55 km E Manalalondo, -19.25583, 47.17751, 1987 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550m, montane rainforest, Malaise trap, 22-26 January 2003 (BLF), collection code: BLF07155, specimen code: CASENT0049853 (CAS). Paratype. One worker. Madagascar, Province Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300m, montane rainforest, pitfall trap, 5-13 December 2000 (BLF), collection code: BLF02370, specimen code: CASENT0409150 (CAS).
Worker diagnosis. Integument black, entirely but finely sculptured, appendages dark brown; scape circular; anterior margin of clypeus rounded, not projected; promesonotal suture impressed, dorsolateral margin of propodeal margin.
Description of minor worker. Medium-sized. Absolute cephalic size (CS: 1.02 ± 0.28; 0.92-1.14). In full-face view, head subquadrate (CWb/CL: 0.39±0.01, 0.39-0.40), posterior margin of head broadly convex, lateral margin of head slightly convex. Eyes sub-elliptical, placed laterally, closer to the posterior margin (PoOC/CL: 0.09±0.01; 0.08-0.09). Mandibles triangular with five teeth. Anterior clypeal margin rounded and not produced anteriorly (ClyL/GPD: 0.24±0.01). Antennal scape virtually circular, surpassing posterior cephalic margin by the length of two basal flagella (SL/ CS: 0.34±0.04; 0.33-0.36). In lateral view, metanotal groove deeply impressed; pronotum, mesonotum, and propodeum dorsum marginated laterally; mesonotum dorsum inclined posteriorly; propodeum dorsum slightly concave, its lateral carina forming a blunt tubercle with the lateral margin of the declivity (MPD/ML: 0.25±0.02; 0.24-0.25). In lateral view, petiolar node strongly convex anterodorsally and flat posteriorly, node summit convex. Head finely areolate and mesosoma feebly areolate-costulate, gastral tergite finely imbricate. Mandible glabrous. Vertex with three pairs of filiform erect setae; mesonotum and propodeum dorsum with single pair of whitish setae; gastral tergite with short, whitish setae aligned transversally on the middle length of each segment; petiolar node with two or three pairs of whitish setae on its dorsum. Body uniformly dark blackish brown to black, appendages brownish, mandible light brown.
Description of major worker. With characteristics exactly the same as minor worker, but head more trapezoidal (CS: 1.74±0.04, 1.71-1.77).
Distribution and biology. Camponotus zoro has been collected by beating low vegetation, sifting leaf litter, and baiting with honey and tuna fish. One worker has been found in Malaise traps from deciduous forest in southern Madagascar (Fig. 72E). In addition, this species has been collected from montane rainforest and Uapaca woodland.
Discussion. Camponotus zoro is characterized by a shallow but distinct metanotal groove, and should not be confused with any other species because of the margination of its mesosoma.
The revision of a large genus such as Camponotus requires the integration of multiple lines of taxonomic evidence to delineate species, namely, original morphological descriptions, detailed quantitative morphological measurements, information on ecological distribution, genetic sequence data, and images of type specimens.