Research Article |
Corresponding author: Jean Claude Rakotonirina ( jcrakoto25@yahoo.com ) Academic editor: Marek Borowiec
© 2022 Jean Claude Rakotonirina, Brian L. Fisher.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rakotonirina JC, Fisher BL (2022) Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology. ZooKeys 1098: 1-180. https://doi.org/10.3897/zookeys.1098.73223
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The Camponotus subgenus Myrmosaga subgen. rev. from the Malagasy region is revised based on analysis of both qualitative morphological characters and morphometric traits. The multivariate analysis used the Nest Centroid (NC)-clustering method to generate species hypotheses based on 19 continuous morphological traits of minor workers. The proposed species hypotheses were confirmed by cumulative Linear Discriminant Analysis (LDA). Morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. The present study places the subgenus Myrmopytia syn. nov. in synonymy to Myrmosaga. It recognizes 38 species, of which 19 are newly described: C. aina sp. nov., C. aro sp. nov., C. asara sp. nov., C. atimo sp. nov., C. bemaheva sp. nov., C. bozaka sp. nov., C. daraina sp. nov., C. harenarum sp. nov., C. joany sp. nov., C. karsti sp. nov., C. kelimaso sp. nov., C. lokobe sp. nov., C. mahafaly sp. nov., C. niavo sp. nov., C. rotrae sp. nov., C. sambiranoensis sp. nov., C. tapia sp. nov., C. tendryi sp. nov., C. vano sp. nov. Eleven species are redescribed: C. aurosus Roger, C. cervicalis Roger, C. dufouri Forel, C. gibber Forel, C. hagensii Forel, C. hova Forel, C. hovahovoides Forel, C. immaculatus Forel, C. quadrimaculatus Forel, C. roeseli Forel, C. strangulatus Santschi. The following are raised to species and redescribed: C. becki Santschi stat. nov., C. boivini Forel stat. rev., C. cemeryi Özdikmen stat. rev., C. mixtellus Forel stat. nov., C. radamae Forel stat. nov. Camponotus maculatus st. fairmairei Santschi syn. nov., is synonymized under C. boivini. The following are synonymized under C. cervicalis: Camponotus cervicalis gaullei Santschi, syn. nov.; Camponotus perroti Forel, syn. nov.; Camponotus perroti aeschylus Forel, syn. nov.; Camponotus gerberti Donisthorpe, syn. nov. Camponotus dufouri imerinensis Forel, syn. nov. is a synonym of C. dufouri, Camponotus hova var. obscuratus Emery, syn. nov. is a synonym of C. hova, Camponotus quadrimaculatus opacata Emery, syn. nov. is a synonym of C. immaculatus, Camponotus maculatus st. legionarium Santschi, syn. nov. is a synonym of C. roeseli, Camponotus hova maculatoides Emery, syn. nov. is a synonym of C. strangulatus. The following are synonymized under C. quadrimaculatus: Camponotus kelleri Forel, syn. nov., Camponotus kelleri var. invalidus Forel, syn. nov., Camponotus quadrimaculatus sellaris Emery, syn. nov. As C. imitator Forel, C. liandia Rakotonirina & Fisher, and C. lubbocki Forel have been recently described and redescribed, only diagnoses and taxonomic discussions are provided. This revision also includes an illustrated species identification key, taxonomic discussions, images, and distribution maps for each species superimposed on the ecoregions of Madagascar.
Madagascar, morphometry, species delimitation, subgenus Myrmosaga, taxonomy
Given the striking morphological variation of Malagasy Camponotus, taxonomic knowledge of this genus has been advanced recently using a combination of traditional morphology-based study and a morphometry-based approach. These techniques and the use of other sources of evidence have helped assign individual specimens to a species, facilitate species recognition, and improve precision of species delimitation in recent revisions (
Myrmosaga was created by
Members of Myrmosaga are found only in Madagascar and surrounding islands of the southwest Indian Ocean region. They occupy all the terrestrial ecoregions encountered in this region. Morphologically, the worker castes of the subgenus present a wide range of features that are the result of its substantial adaptive radiation across these islands, but particularly in Madagascar. Several species within the subgenus show development of a few or many characters similar to fauna from the Afrotropical region. Our understanding of this observed morphological diversification has triggered taxonomic changes for some of the species that were described and combined previously under different subgenera of Camponotus (Table
Summary of the new subgenus placements for the Malagasy Camponotus (Myrmosaga).
Taxon | Changes | Old subgenus placement | New subgenus placement |
---|---|---|---|
Myrmopytia | syn. nov. | Myrmosaga | |
Camponotus aurosus | subgenus | Myrmosericus | Myrmosaga |
Camponotus becki | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus boivini | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus cemeryi | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus cervicalis | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus dufouri | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus gibber | subgenus | Mayria | Myrmosaga |
Camponotus gouldi | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus hagensii | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus hova | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus hovahovoides | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus imitator | subgenus | Myrmopytia | Myrmosaga |
Camponotus immaculatus | subgenus | Mayria | Myrmosaga |
Camponotus liandia | subgenus | Mayria | Myrmosaga |
Camponotus lubbocki | subgenus | Mayria | Myrmosaga |
Camponotus mixtellus | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus quadrimaculatus | subgenus | Mayria | Myrmosaga |
Camponotus radamae | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus roeseli | subgenus | Tanaemyrmex | Myrmosaga |
Camponotus strangulatus | subgenus | Tanaemyrmex | Myrmosaga |
This proposed Myrmosaga classification is based on the use of additional morphological features thought to be relevant in defining the subgenus more precisely. Minor workers of Myrmosaga are generally characterized by the combination of the following morphological characters: clypeus medially carinate, mandible armed with six teeth, laterodorsal angle of mesosoma never marginate, petiolar node never conical.
The present revision recognizes 38 species in the subgenus Myrmosaga, provides an illustrated species-level identification key, and a description of each species complemented by high-resolution montage images and a geographic distribution map.
PSWC P.S. Ward Collection, University of California at Davis, CA, USA.
ZMHB Museum für Naturkunde der Humboldt Universität, Berlin, Germany.
The present study includes all specimens of the subgenus Myrmosaga collected from the arthropod survey project conducted in Madagascar and surrounding islands in the Malagasy region by the members of the Madagascar Biodiversity Center and other ant researchers. Most of these collections were sampled from 1992 through 2018. All pinned specimens examined in this study are available on the AntWeb portal (http://www.antweb.org) and can be accessed using the unique identifying specimen code (CASENTnumber) assigned to each pinned specimen.
A total of 776 specimens from 324 collecting events was measured in this study (see Suppl. material
Using a Leica MZ12 binocular microscope, qualitative morphological examinations were conducted on minor and major workers to evaluate patterns of morphological discontinuities and phenotypic similarity. Observed discontinuities in morphological space could indicate reproductive boundaries between populations and as such were used to infer potential species limits. Species limits based on qualitative morphological characters were compared with the species hypothesis derived from the quantitative analysis of measurements. Differences between the sizes of the measured worker castes in the present study might result in a conflict between the two approaches. To avoid this conflict any specimen with unusual size class was removed from the analysis.
Digital color images of lateral and dorsal views of the entire body and full-face views of the head of each species were created using a JVC KY-75 or a Leica DFC450 digital camera with a Leica Z16 APO microscope and Leica Application Suite software (v3.8). These images are also available online on AntWeb (www.antweb.org) and are accessible using the unique identifying specimen code.
Distribution maps for all species were generated by importing specimen distribution records into the Diva GIS program (
Article 74 in the ICZN’s code states that a lectotype may be designated from syntype specimens which directly match the original description of a named species in order to stabilize the nomenclature. Therefore, the phrase “by present designation” is used to indicate a lectotype. As stated in Article 75.1 of the ICZN’s code, neotype designation is necessary for a name with no extant name-bearing type to objectively define the nominal taxon. Thus, a neotype has been selected according to the qualifying conditions specified in Article 75.3.
Etymologies are provided for new species to facilitate name stability. Etymologies indicate if the new name is a non-Latin or Latin (or Latinized) word. For Latin names, we also include the part of speech (nouns, adjectives, participles), gender (masculine, feminine, or neuter), number (singular or plural) and grammatical case (e.g. subjective/nominative, possessive/genitive).
Morphometric measurements of the minor and major workers were taken using a Leica MZ12 stereomicroscope equipped with a cross-scaled ocular micrometer and an orthogonal pair of micrometers. All measurements and indices are presented as arithmetic means and ranges are shown as minimum and maximum values in parentheses. Body size dimensions are expressed in millimeters (mm) and all values were rounded to the second decimal place.
The following 19 morphometric measurements were evaluated in the present revision:
CL (Maximum cephalic length): The maximum midline length of the head in full-face view, measured from the midpoint of the posterior margin to the midpoint of the anterior margin of the clypeus.
ClyL (Clypeal length): the maximum midline length of the clypeus measured from the posterior margin to the anterior margin in anterodorsal view, in which the anterior and posterior clypeal margins are aligned to the same focus. Median concavity on either or both margins reduces the length of the clypeus.
CS (Cephalic size): The arithmetic mean of CL and CWb. CS is used to indicate the general body size of the ant.
CW (Maximum cephalic width): The maximum distance between the lateral margins of the compound eyes in full-face view.
CWb (Maximum head capsule width): The maximum width of the head excluding the compound eyes.
EL (Eye length): Maximum diameter of the compound eye.
FR (Frontal carina distance): The maximum distance between the frontal carinae.
GPD (Maximum tentorial pit distance): The longest distance between the centers of the fossae located at or very close to the posterolateral margin of the clypeus.
HTL (Maximum hind tibia length): Straight line length of the hind tibia measured from the constriction immediately before its proximal insertion to its distalmost point, excluding the bristles or spines. The measurement of the tibia is taken from any view where the constriction before its proximal insertion is visible.
ML (Mesosoma length): The longest median anatomical line that connects the posteriormost point of the propodeal lobe with the anteriormost point of the pronotal collar; preferentially measured in lateral view, but if one of the reference points is not visible, dorsal view may be used.
MPD (Mesothoracico-propodeal distance): With the promesonotal suture and the anterior petiolar foramen margin in the same plane of focus in dorsal view, the maximum midline length between the promesonotal suture and the posteriormost point of the propodeal process dorsal to the petiolar insertion (see
MPH (Mesothoracico-propodeal height): With the mesosoma in lateral view, the length of the line between the anteroventral corner of the mesopleuron dorsal to the insertion of the mesocoxa, and the dorsalmost point of the propodeum that is crossed by the measured line. The line is perpendicular to the diagonal line of the mesosoma that connects the anteriormost point of the pronotal shield and the posteriormost point of the propodeal process dorsal to the petiolar insertion, in lateral view.
MW (Mesosoma width): Maximum width of the pronotum in dorsal view, which in the subgenus Myrmosaga is also the maximum mesosomal width (hence “mesosoma width”).
NOH (Petiolar node height): The maximum distance between the petiolar spiracle and the dorsalmost point of the petiolar node.
OMD (Oculo-mandibular distance): The smallest distance between the anterior margin of the compound eye and the mandibular insertion to the head.
PEW (Petiolar width): The maximum width of the petiole in dorsal view.
PoOc (Postocular distance): The distance between the posteromedian margin of the head and the level of the posterior margin of the compound eyes measured along the midline of the head in full-face view.
PrOc (Preocular distance): The distance between the anteromedian margin of the clypeus and the level of the anterior margin of the compound eyes measured along the midline of the head in full-face view.
SL (Scape length): Straight line length of the first antennal segment excluding the basal condyle.
TCD (Torular carina distance): The minimum distance between the torular arches that surround the antennal insertion.
The datasets assessed in the present study consist of (1) the primary measurement data of the 19 morphological characters and the one calculated character CS (a widely applied size indicator) of each measured specimen (see table of basic measurements of the specimens in the Suppl. material
Ratios of morphometric data for minors and majors of the species. Upper line: mean of ratios ± standard deviation, lower line in square brackets: minimum and maximum values. Note: if two or more specimens were available, then minimum, maximum values are given; one value is presented if only one specimen was available.
Species | Castes | CS | CWb/CL | CW/CL | PrOc/CL | PoOc/CL | FR/CS | TCD/CS | ClyL/CL | ClyL/GPD | SL/CS | EL/CS | OMD/CS | MW/ML | PEW/CS | MPD/ML | HTL/CS | ML/CS | MPH/ML | NOH/CS |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
aina | Minor (n= 2) | 4.41±0.74 | 0.72±0.00 | 0.69±0.01 | 0.49±0.01 | 0.26±0.00 | 0.28±0.01 | 0.21±0.01 | 0.29±0.01 | 0.74±0.07 | 1.64±0.02 | 0.24±0.02 | 0.47±0.03 | 0.38±0.01 | 0.21±0.04 | 0.66±0.05 | 1.94±0.04 | 1.91±0.03 | 0.31±0.09 | 0.17±0.01 |
[3.89, 4.94] | [0.72, 0.72] | [0.68, 0.70] | [0.49, 0.50] | [0.26, 0.26] | [0.27, 0.29] | [0.20, 0.21] | [0.28, 0.29] | [0.69, 0.80] | [1.62, 1.66] | [0.22, 0.26] | [0.46, 0.49] | [0.37, 0.39] | [0.19, 0.24] | [0.63, 0.69] | [1.91, 1.97] | [1.89, 1.93] | [0.25, 0.37] | [0.16, 0.18] | ||
aro | Minor (n= 31) | 1.78±0.15 | 0.72±0.05 | 0.67±0.04 | 0.50±0.01 | 0.28±0.01 | 0.25±0.01 | 0.23±0.01 | 0.28±0.01 | 0.72±0.04 | 1.59±0.08 | 0.25±0.01 | 0.53±0.02 | 0.33±0.01 | 0.22±0.01 | 0.68±0.01 | 1.96±0.09 | 1.86±0.06 | 0.28±0.01 | 0.23±0.01 |
[1.56, 2.09] | [0.62, 0.79] | [0.57, 0.70] | [0.48, 0.53] | [0.27, 0.30] | [0.24, 0.27] | [0.22, 0.25] | [0.24, 0.29] | [0.63, 0.77] | [1.46, 1.71] | [0.23, 0.26] | [0.52, 0.58] | [0.32, 0.35] | [0.20, 0.24] | [0.66, 0.71] | [1.76, 2.08] | [1.73, 1.94] | [0.26, 0.30] | [0.21, 0.24] | ||
Major (n= 5) | 3.10±0.18 | 0.92±0.01 | 0.75±0.01 | 0.55±0.01 | 0.27±0.01 | 0.25±0.00 | 0.22±0.00 | 0.30±0.00 | 0.82±0.01 | 0.99±0.06 | 0.20±0.01 | 0.51±0.01 | 0.59±0.01 | 0.19±0.01 | 0.49±0.01 | 1.33±0.06 | 0.93±0.04 | 1.47±0.02 | 0.18±0.00 | |
[2.82, 3.29] | [0.90, 0.94] | [0.74, 0.77] | [0.54, 0.56] | [0.25, 0.28] | [0.24, 0.25] | [0.22, 0.23] | [0.29, 0.30] | [0.81, 0.83] | [0.93, 1.07] | [0.19, 0.21] | [0.50, 0.52] | [0.57, 0.60] | [0.19, 0.20] | [0.48, 0.50] | [1.26, 1.41] | [0.91, 1.00] | [1.44, 1.49] | [0.18, 0.19] | ||
asara | Minor (n= 19) | 1.67±0.13 | 0.72±0.07 | 0.67±0.01 | 0.51±0.02 | 0.26±0.01 | 0.26±0.01 | 0.22±0.01 | 0.29±0.01 | 0.72±0.02 | 1.57±0.08 | 0.26±0.01 | 0.52±0.02 | 0.34±0.01 | 0.25±0.01 | 0.71±0.01 | 1.89±0.09 | 1.86±0.07 | 0.34±0.01 | 0.26±0.02 |
[1.49, 2.02] | [0.67, 0.97] | [0.65, 0.70] | [0.49, 0.54] | [0.24, 0.28] | [0.23, 0.27] | [0.19, 0.23] | [0.28, 0.30] | [0.68, 0.76] | [1.38, 1.68] | [0.23, 0.28] | [0.44, 0.55] | [0.33, 0.36] | [0.22, 0.26] | [0.69, 0.73] | [1.67, 1.99] | [1.68, 1.95] | [0.33, 0.38] | [0.23, 0.29] | ||
Major (n= 4) | 3.22±0.15 | 0.93±0.01 | 0.77±0.01 | 0.54±0.01 | 0.27±0.01 | 0.24±0.03 | 0.21±0.02 | 0.30±0.01 | 0.86±0.03 | 0.90±0.11 | 0.20±0.01 | 0.49±0.01 | 0.58±0.01 | 0.22±0.02 | 0.53±0.02 | 1.23±0.05 | 0.98±0.03 | 1.38±0.03 | 0.19±0.01 | |
[3.03, 3.39] | [0.92, 0.93] | [0.75, 0.78] | [0.53, 0.55] | [0.27, 0.28] | [0.19, 0.26] | [0.18, 0.22] | [0.30, 0.31] | [0.83, 0.89] | [0.74, 0.99] | [0.19, 0.20] | [0.48, 0.49] | [0.56, 0.59] | [0.21, 0.25] | [0.51, 0.55] | [1.18, 1.29] | [0.95, 1.02] | [1.33, 1.41] | [0.18, 0.21] | ||
atimo | Minor (n= 31) | 1.56±0.12 | 0.66±0.03 | 0.66±0.02 | 0.50±0.01 | 0.27±0.01 | 0.23±0.01 | 0.21±0.01 | 0.28±0.01 | 0.74±0.03 | 1.66±0.09 | 0.28±0.01 | 0.51±0.01 | 0.34±0.02 | 0.22±0.01 | 0.69±0.01 | 1.89±0.09 | 1.97±0.06 | 0.33±0.02 | 0.24±0.02 |
[1.28, 1.79] | [0.59, 0.72] | [0.62, 0.68] | [0.47, 0.52] | [0.25, 0.29] | [0.22, 0.25] | [0.20, 0.23] | [0.27, 0.29] | [0.67, 0.81] | [1.48, 1.84] | [0.26, 0.30] | [0.48, 0.53] | [0.30, 0.37] | [0.21, 0.25] | [0.65, 0.71] | [1.72, 2.08] | [1.85, 2.11] | [0.29, 0.36] | [0.20, 0.27] | ||
Major (n= 8) | 3.23±0.34 | 0.97±0.04 | 0.77±0.02 | 0.53±0.01 | 0.29±0.01 | 0.24±0.01 | 0.21±0.00 | 0.29±0.01 | 0.86±0.03 | 0.89±0.06 | 0.19±0.01 | 0.47±0.01 | 0.59±0.02 | 0.19±0.01 | 0.55±0.03 | 1.09±0.05 | 0.93±0.05 | 1.41±0.05 | 0.19±0.01 | |
[2.80, 3.77] | [0.89, 1.03] | [0.73, 0.79] | [0.52, 0.55] | [0.27, 0.30] | [0.23, 0.24] | [0.20, 0.21] | [0.28, 0.30] | [0.82, 0.89] | [0.80, 1.00] | [0.17, 0.20] | [0.46, 0.48] | [0.56, 0.63] | [0.18, 0.21] | [0.52, 0.62] | [1.02, 1.20] | [0.84, 1.00] | [1.36, 1.51] | [0.17, 0.21] | ||
aurosus | Minor (n= 16) | 1.37±0.05 | 0.74±0.02 | 0.74±0.01 | 0.54±0.01 | 0.24±0.01 | 0.30±0.01 | 0.25±0.01 | 0.33±0.01 | 0.79±0.02 | 1.33±0.04 | 0.27±0.01 | 0.53±0.01 | 0.39±0.02 | 0.25±0.02 | 0.73±0.04 | 1.48±0.05 | 1.74±0.08 | 0.36±0.03 | 0.25±0.03 |
[1.27, 1.45] | [0.71, 0.77] | [0.72, 0.76] | [0.52, 0.56] | [0.22, 0.26] | [0.29, 0.32] | [0.24, 0.26] | [0.32, 0.34] | [0.76, 0.85] | [1.27, 1.40] | [0.25, 0.29] | [0.51, 0.54] | [0.34, 0.41] | [0.22, 0.29] | [0.61, 0.76] | [1.39, 1.54] | [1.67, 2.04] | [0.30, 0.41] | [0.22, 0.30] | ||
Major (n= 4) | 2.24±0.11 | 0.92±0.03 | 0.81±0.02 | 0.57±0.01 | 0.24±0.01 | 0.29±0.01 | 0.26±0.00 | 0.35±0.01 | 0.92±0.02 | 0.93±0.06 | 0.21±0.02 | 0.49±0.01 | 0.60±0.02 | 0.21±0.01 | 0.54±0.02 | 1.02±0.08 | 0.95±0.05 | 1.38±0.02 | 0.23±0.08 | |
[2.15, 2.37] | [0.87, 0.95] | [0.79, 0.82] | [0.56, 0.59] | [0.22, 0.25] | [0.28, 0.30] | [0.25, 0.26] | [0.34, 0.36] | [0.90, 0.94] | [0.86, 0.99] | [0.19, 0.23] | [0.47, 0.50] | [0.59, 0.64] | [0.20, 0.21] | [0.50, 0.56] | [0.94, 1.12] | [0.90, 1.00] | [1.37, 1.41] | [0.17, 0.35] | ||
becki | Minor (n= 11) | 1.63±0.18 | 0.77±0.03 | 0.75±0.01 | 0.50±0.01 | 0.26±0.01 | 0.27±0.01 | 0.24±0.01 | 0.30±0.01 | 0.72±0.03 | 1.27±0.05 | 0.27±0.02 | 0.49±0.01 | 0.38±0.02 | 0.22±0.01 | 0.72±0.03 | 1.47±0.05 | 1.72±0.09 | 0.37±0.02 | 0.24±0.06 |
[1.34, 1.96] | [0.73, 0.83] | [0.73, 0.77] | [0.48, 0.51] | [0.24, 0.27] | [0.26, 0.29] | [0.22, 0.25] | [0.29, 0.31] | [0.67, 0.77] | [1.15, 1.34] | [0.25, 0.30] | [0.48, 0.50] | [0.35, 0.43] | [0.21, 0.24] | [0.70, 0.78] | [1.38, 1.56] | [1.58, 1.91] | [0.33, 0.42] | [0.2, 0.28] | ||
Major (n= 3) | 2.80±0.23 | 0.97±0.03 | 0.83±0.02 | 0.53±0.01 | 0.27±0.01 | 0.26±0.00 | 0.22±0.00 | 0.32±0.01 | 0.85±0.01 | 0.84±0.03 | 0.21±0.01 | 0.46±0.01 | 0.87±0.43 | 0.19±0.01 | 0.55±0.01 | 1.03±0.03 | 0.95±0.02 | 1.39±0.02 | 0.20±0.02 | |
[2.54, 2.96] | [0.94, 1.00] | [0.80, 0.84] | [0.51, 0.54] | [0.26, 0.27] | [0.26, 0.26] | [0.22, 0.22] | [0.31, 0.33] | [0.84, 0.87] | [0.81, 0.87] | [0.20, 0.22] | [0.46, 0.47] | [0.62, 1.37] | [0.18, 0.19] | [0.54, 0.56] | [1.00, 1.06] | [0.93, 0.96] | [1.38, 1.41] | [0.18, 0.22] | ||
bemaheva | Minor (n= 23) | 1.63±0.13 | 0.64±0.03 | 0.64±0.01 | 0.49±0.01 | 0.26±0.01 | 0.26±0.01 | 0.22±0.01 | 0.29±0.01 | 0.70±0.02 | 1.63±0.10 | 0.30±0.01 | 0.50±0.02 | 0.37±0.01 | 0.28±0.02 | 0.75±0.01 | 1.80±0.08 | 2.02±0.07 | 0.40±0.02 | 0.28±0.02 |
[1.28, 1.79] | [0.53, 0.68] | [0.62, 0.67] | [0.47, 0.50] | [0.24, 0.28] | [0.25, 0.29] | [0.20, 0.24] | [0.27, 0.29] | [0.66, 0.77] | [1.47, 1.87] | [0.29, 0.34] | [0.48, 0.54] | [0.35, 0.39] | [0.24, 0.31] | [0.72, 0.77] | [1.64, 2.03] | [1.95, 2.24] | [0.37, 0.43] | [0.24, 0.31] | ||
boivini | Minor (n= 49) | 1.15±0.06 | 0.67±0.03 | 0.71±0.03 | 0.46±0.01 | 0.27±0.01 | 0.32±0.01 | 0.25±0.01 | 0.29±0.01 | 0.72±0.04 | 1.37±0.10 | 0.32±0.02 | 0.47±0.02 | 0.38±0.01 | 0.27±0.02 | 0.75±0.01 | 1.57±0.12 | 1.83±0.07 | 0.39±0.01 | 0.26±0.02 |
[1.00, 1.27] | [0.61, 0.73] | [0.65, 0.77] | [0.43, 0.49] | [0.24, 0.30] | [0.30, 0.35] | [0.23, 0.26] | [0.27, 0.31] | [0.63, 0.79] | [1.22, 1.56] | [0.29, 0.36] | [0.44, 0.51] | [0.34, 0.40] | [0.22, 0.30] | [0.71, 0.78] | [1.41, 1.79] | [1.70, 2.00] | [0.37, 0.41] | [0.23, 0.30] | ||
Major (n= 13) | 2.29±0.28 | 0.91±0.04 | 0.79±0.02 | 0.51±0.01 | 0.29±0.01 | 0.33±0.01 | 0.25±0.01 | 0.32±0.01 | 0.89±0.02 | 0.71±0.06 | 0.22±0.02 | 0.44±0.01 | 0.59±0.01 | 0.22±0.01 | 0.57±0.02 | 0.94±0.07 | 0.93±0.05 | 1.33±0.02 | 0.20±0.02 | |
[1.96, 3.12] | [0.86, 0.99] | [0.77, 0.83] | [0.49, 0.52] | [0.27, 0.33] | [0.32, 0.35] | [0.24, 0.26] | [0.30, 0.34] | [0.85, 0.93] | [0.58, 0.79] | [0.17, 0.24] | [0.42, 0.45] | [0.57, 0.62] | [0.20, 0.25] | [0.54, 0.60] | [0.75, 1.03] | [0.82, 1.00] | [1.29, 1.38] | [0.17, 0.22] | ||
bozaka | Minor (n= 12) | 1.33±0.13 | 0.77±0.03 | 0.78±0.02 | 0.51±0.01 | 0.25±0.01 | 0.28±0.01 | 0.24±0.01 | 0.30±0.01 | 0.74±0.03 | 1.19±0.05 | 0.28±0.01 | 0.46±0.01 | 0.40±0.01 | 0.27±0.01 | 0.74±0.02 | 1.31±0.06 | 1.67±0.07 | 0.40±0.01 | 0.29±0.02 |
[1.16, 1.59] | [0.73, 0.84] | [0.76, 0.81] | [0.47, 0.53] | [0.23, 0.26] | [0.28, 0.29] | [0.23, 0.29] | [0.29, 0.32] | [0.70, 0.82] | [1.08, 1.25] | [0.26, 0.30] | [0.44, 0.47] | [0.38, 0.42] | [0.26, 0.28] | [0.71, 0.77] | [1.21, 1.38] | [1.55, 1.77] | [0.38, 0.43] | [0.27, 0.32] | ||
Major (n= 6) | 2.07±0.18 | 0.95±0.03 | 0.86±0.01 | 0.51±0.02 | 0.27±0.01 | 0.28±0.01 | 0.24±0.01 | 0.32±0.01 | 0.85±0.02 | 0.90±0.04 | 0.23±0.01 | 0.44±0.01 | 0.58±0.02 | 0.24±0.01 | 0.57±0.01 | 1.02±0.04 | 1.02±0.04 | 1.33±0.02 | 0.25±0.02 | |
[1.91, 2.40] | [0.90, 0.98] | [0.85, 0.88] | [0.49, 0.53] | [0.25, 0.29] | [0.28, 0.30] | [0.23, 0.25] | [0.31, 0.33] | [0.83, 0.88] | [0.86, 0.94] | [0.21, 0.25] | [0.43, 0.45] | [0.56, 0.61] | [0.23, 0.25] | [0.55, 0.58] | [0.98, 1.08] | [0.96, 1.08] | [1.31, 1.38] | [0.22, 0.26] | ||
cemeryi | Minor (n= 9) | 1.11±0.07 | 0.67±0.02 | 0.72±0.02 | 0.47±0.01 | 0.25±0.02 | 0.28±0.01 | 0.24±0.01 | 0.27±0.01 | 0.66±0.03 | 1.49±0.05 | 0.33±0.01 | 0.48±0.01 | 0.41±0.02 | 0.29±0.02 | 0.76±0.02 | 1.58±0.06 | 1.88±0.06 | 0.42±0.02 | 0.27±0.03 |
[0.98, 1.28] | [0.64, 0.71] | [0.67, 0.75] | [0.45, 0.49] | [0.22, 0.29] | [0.26, 0.30] | [0.22, 0.25] | [0.26, 0.29] | [0.62, 0.72] | [1.41, 1.60] | [0.31, 0.35] | [0.47, 0.50] | [0.38, 0.43] | [0.27, 0.34] | [0.72, 0.79] | [1.48, 1.76] | [1.77, 2.01] | [0.39, 0.46] | [0.21, 0.31] | ||
Major (n= 7) | 2.41±0.22 | 0.87±0.05 | 0.75±0.03 | 0.51±0.01 | 0.29±0.01 | 0.28±0.01 | 0.22±0.01 | 0.30±0.01 | 0.86±0.03 | 0.80±0.07 | 0.21±0.01 | 0.46±0.02 | 0.59±0.02 | 0.23±0.01 | 0.58±0.01 | 0.99±0.06 | 0.97±0.04 | 1.32±0.03 | 0.21±0.02 | |
[2.19, 2.72] | [0.83, 0.94] | [0.71, 0.79] | [0.50, 0.52] | [0.27, 0.30] | [0.26, 0.30] | [0.21, 0.23] | [0.29, 0.31] | [0.83, 0.90] | [0.70, 0.91] | [0.19, 0.23] | [0.43, 0.48] | [0.57, 0.63] | [0.22, 0.25] | [0.56, 0.61] | [0.86, 1.06] | [0.92, 1.02] | [1.29, 1.35] | [0.17, 0.23] | ||
cervicalis | Minor (n= 8) | 1.94±0.15 | 0.59±0.03 | 0.58±0.01 | 0.47±0.02 | 0.30±0.01 | 0.27±0.01 | 0.23±0.01 | 0.30±0.01 | 0.81±0.02 | 1.81±0.12 | 0.28±0.01 | 0.54±0.01 | 0.34±0.01 | 0.27±0.01 | 0.71±0.01 | 2.15±0.12 | 2.08±0.08 | 0.36±0.02 | 0.29±0.02 |
[1.83, 2.18] | [0.56, 0.65] | [0.56, 0.60] | [0.45, 0.49] | [0.29, 0.31] | [0.26, 0.28] | [0.22, 0.24] | [0.28, 0.31] | [0.79, 0.85] | [1.57, 1.94] | [0.26, 0.30] | [0.52, 0.56] | [0.32, 0.36] | [0.25, 0.29] | [0.69, 0.73] | [1.91, 2.33] | [1.96, 2.15] | [0.34, 0.38] | [0.27, 0.32] | ||
Major (n= 5) | 4.17±0.35 | 0.91±0.03 | 0.70±0.01 | 0.53±0.01 | 0.28±0.01 | 0.25±0.01 | 0.21±0.01 | 0.33±0.01 | 0.98±0.03 | 0.95±0.06 | 0.19±0.01 | 0.47±0.01 | 0.56±0.02 | 0.22±0.01 | 0.52±0.01 | 1.23±0.07 | 1.00±0.04 | 1.35±0.02 | 0.21±0.01 | |
[3.83, 4.63] | [0.85, 0.93] | [0.68, 0.71] | [0.52, 0.54] | [0.28, 0.29] | [0.24, 0.27] | [0.20, 0.23] | [0.31, 0.33] | [0.94, 1.01] | [0.89, 1.04] | [0.18, 0.21] | [0.46, 0.49] | [0.53, 0.58] | [0.21, 0.23] | [0.51, 0.53] | [1.18, 1.35] | [0.97, 1.07] | [1.32, 1.36] | [0.19, 0.22] | ||
daraina | Minor (n= 19) | 1.87±0.17 | 0.60±0.02 | 0.59±0.01 | 0.48±0.01 | 0.29±0.01 | 0.25±0.00 | 0.22±0.01 | 0.28±0.01 | 0.75±0.02 | 1.72±0.06 | 0.28±0.01 | 0.52±0.01 | 0.34±0.01 | 0.28±0.01 | 0.72±0.01 | 2.04±0.05 | 2.09±0.04 | 0.37±0.01 | 0.30±0.02 |
[1.68, 2.31] | [0.57, 0.66] | [0.57, 0.62] | [0.45, 0.50] | [0.28, 0.30] | [0.24, 0.26] | [0.21, 0.23] | [0.27, 0.30] | [0.71, 0.79] | [1.56, 1.82] | [0.26, 0.30] | [0.50, 0.52] | [0.32, 0.36] | [0.26, 0.31] | [0.71, 0.74] | [1.93, 2.15] | [1.95, 2.16] | [0.35, 0.38] | [0.26, 0.32] | ||
Major (n= 5) | 3.54±0.25 | 0.90±0.04 | 0.72±0.02 | 0.53±0.01 | 0.28±0.01 | 0.25±0.01 | 0.21±0.01 | 0.30±0.01 | 0.86±0.03 | 0.96±0.06 | 0.20±0.01 | 0.47±0.01 | 0.56±0.01 | 0.22±0.01 | 0.54±0.01 | 1.23±0.06 | 1.03±0.03 | 1.35±0.03 | 0.20±0.01 | |
[3.12, 3.76] | [0.85, 0.94] | [0.70, 0.75] | [0.52, 0.55] | [0.28, 0.29] | [0.24, 0.26] | [0.20, 0.21] | [0.30, 0.31] | [0.82, 0.89] | [0.90, 1.04] | [0.18, 0.22] | [0.46, 0.48] | [0.54, 0.58] | [0.21, 0.24] | [0.53, 0.54] | [1.18, 1.31] | [1.00, 1.07] | [1.32, 1.40] | [0.19, 0.21] | ||
dufouri | Minor (n= 19) | 1.97±0.19 | 0.56±0.02 | 0.55±0.02 | 0.45±0.01 | 0.33±0.01 | 0.25±0.01 | 0.21±0.01 | 0.27±0.01 | 0.82±0.03 | 1.93±0.08 | 0.28±0.02 | 0.51±0.01 | 0.28±0.01 | 0.24±0.01 | 0.67±0.01 | 2.37±0.09 | 2.15±0.08 | 0.29±0.01 | 0.29±0.02 |
[1.66, 2.26] | [0.53, 0.59] | [0.53, 0.59] | [0.43, 0.47] | [0.31, 0.36] | [0.24, 0.27] | [0.19, 0.22] | [0.26, 0.28] | [0.77, 0.86] | [1.76, 2.12] | [0.25, 0.31] | [0.49, 0.52] | [0.27, 0.32] | [0.22, 0.27] | [0.65, 0.71] | [2.23, 2.57] | [1.98, 2.31] | [0.27, 0.32] | [0.26, 0.33] | ||
Major (n= 5) | 3.93±0.51 | 0.88±0.08 | 0.68±0.03 | 0.51±0.02 | 0.32±0.01 | 0.24±0.01 | 0.20±0.01 | 0.30±0.01 | 0.94±0.02 | 1.02±0.08 | 0.19±0.01 | 0.47±0.01 | 0.51±0.02 | 0.18±0.01 | 0.46±0.02 | 1.34±0.13 | 0.99±0.06 | 1.38±0.04 | 0.19±0.02 | |
[3.46, 4.54] | [0.80, 0.97] | [0.65, 0.72] | [0.49, 0.53] | [0.30, 0.33] | [0.23, 0.25] | [0.19, 0.21] | [0.29, 0.31] | [0.93, 0.98] | [0.94, 1.10] | [0.18, 0.20] | [0.46, 0.47] | [0.49, 0.54] | [0.17, 0.19] | [0.44, 0.48] | [1.18, 1.46] | [0.92, 1.06] | [1.33, 1.41] | [0.18, 0.22] | ||
gibber | Minor (n= 37) | 1.24±0.13 | 0.85±0.03 | 0.83±0.02 | 0.50±0.01 | 0.25±0.01 | 0.35±0.01 | 0.28±0.01 | 0.31±0.01 | 0.69±0.04 | 1.17±0.07 | 0.27±0.01 | 0.46±0.01 | 0.44±0.02 | 0.29±0.01 | 0.74±0.02 | 1.20±0.07 | 1.57±0.07 | 0.36±0.02 | 0.21±0.02 |
[1.00, 1.51] | [0.79, 0.93] | [0.79, 0.89] | [0.47, 0.54] | [0.23, 0.28] | [0.33, 0.36] | [0.26, 0.29] | [0.29, 0.33] | [0.62, 0.76] | [0.93, 1.27] | [0.24, 0.30] | [0.44, 0.49] | [0.40, 0.50] | [0.27, 0.32] | [0.71, 0.79] | [1.00, 1.30] | [1.38, 1.67] | [0.32, 0.41] | [0.17, 0.24] | ||
Major (n= 9) | 2.45±0.21 | 1.05±0.04 | 0.90±0.04 | 0.52±0.03 | 0.28±0.02 | 0.33±0.02 | 0.26±0.01 | 0.33±0.01 | 0.83±0.03 | 0.74±0.06 | 0.21±0.01 | 0.45±0.02 | 0.66±0.02 | 0.25±0.02 | 0.54±0.01 | 0.86±0.05 | 0.88±0.04 | 1.32±0.03 | 0.17±0.01 | |
[2.21, 2.82] | [1.01, 1.10] | [0.86, 0.97] | [0.50, 0.60] | [0.26, 0.31] | [0.31, 0.36] | [0.24, 0.29] | [0.31, 0.36] | [0.76, 0.86] | [0.68, 0.85] | [0.19, 0.22] | [0.43, 0.50] | [0.63, 0.70] | [0.22, 0.27] | [0.52, 0.57] | [0.81, 0.98] | [0.84, 0.97] | [1.28, 1.37] | [0.15, 0.19] | ||
gouldi | Minor (n= 18) | 2.52±0.17 | 0.66±0.02 | 0.62±0.01 | 0.50±0.01 | 0.29±0.02 | 0.26±0.01 | 0.23±0.01 | 0.29±0.01 | 0.77±0.11 | 1.56±0.06 | 0.24±0.01 | 0.49±0.01 | 0.47±0.01 | 0.25±0.01 | 0.49±0.02 | 2.11±0.08 | 1.47±0.03 | 1.36±0.03 | 0.29±0.02 |
[2.29, 2.89] | [0.64, 0.70] | [0.60, 0.65] | [0.49, 0.55] | [0.26, 0.32] | [0.25, 0.27] | [0.21, 0.24] | [0.27, 0.31] | [0.68, 1.16] | [1.44, 1.66] | [0.23, 0.26] | [0.47, 0.50] | [0.44, 0.48] | [0.23, 0.27] | [0.47, 0.53] | [1.97, 2.26] | [1.42, 1.51] | [1.31, 1.41] | [0.24, 0.31] | ||
Major (n= 6) | 5.19±0.23 | 0.98±0.01 | 0.76±0.01 | 0.55±0.01 | 0.28±0.01 | 0.24±0.01 | 0.21±0.00 | 0.33±0.01 | 0.93±0.02 | 0.89±0.05 | 0.17±0.01 | 0.47±0.01 | 0.57±0.01 | 0.20±0.01 | 0.53±0.02 | 1.19±0.05 | 0.96±0.02 | 1.34±0.03 | 0.23±0.02 | |
[4.89, 5.51] | [0.97, 1.00] | [0.74, 0.76] | [0.54, 0.55] | [0.27, 0.30] | [0.23, 0.25] | [0.20, 0.21] | [0.32, 0.34] | [0.90, 0.95] | [0.84, 0.95] | [0.16, 0.18] | [0.46, 0.48] | [0.57, 0.58] | [0.19, 0.21] | [0.50, 0.55] | [1.16, 1.26] | [0.93, 0.99] | [1.31, 1.38] | [0.20, 0.24] | ||
hagensii | Minor (n= 17) | 1.45±0.14 | 0.80±0.03 | 0.80±0.02 | 0.48±0.01 | 0.25±0.01 | 0.34±0.01 | 0.27±0.01 | 0.31±0.01 | 0.74±0.04 | 1.24±0.06 | 0.30±0.01 | 0.46±0.01 | 0.41±0.01 | 0.30±0.01 | 0.75±0.01 | 1.37±0.06 | 1.66±0.04 | 0.38±0.02 | 0.24±0.02 |
[1.30, 1.68] | [0.74, 0.84] | [0.76, 0.83] | [0.46, 0.50] | [0.23, 0.25] | [0.31, 0.35] | [0.26, 0.29] | [0.30, 0.34] | [0.68, 0.85] | [1.09, 1.36] | [0.28, 0.33] | [0.43, 0.48] | [0.39, 0.42] | [0.27, 0.32] | [0.73, 0.77] | [1.23, 1.48] | [1.56, 1.74] | [0.36, 0.41] | [0.21, 0.27] | ||
Major (n= 5) | 2.37±0.26 | 0.97±0.05 | 0.87±0.03 | 0.50±0.01 | 0.26±0.02 | 0.33±0.01 | 0.27±0.01 | 0.34±0.01 | 0.86±0.02 | 0.86±0.09 | 0.24±0.02 | 0.45±0.01 | 0.61±0.02 | 0.25±0.02 | 0.56±0.02 | 1.03±0.09 | 0.98±0.06 | 1.33±0.04 | 0.19±0.01 | |
[2.00, 2.70] | [0.92, 1.04] | [0.85, 0.91] | [0.50, 0.51] | [0.24, 0.28] | [0.31, 0.35] | [0.25, 0.27] | [0.33, 0.34] | [0.84, 0.90] | [0.74, 0.97] | [0.22, 0.27] | [0.44, 0.47] | [0.58, 0.64] | [0.22, 0.26] | [0.54, 0.58] | [0.90, 1.12] | [0.92, 1.07] | [1.27, 1.38] | [0.18, 0.21] | ||
harenarum | Minor (n= 5) | 1.53±0.11 | 0.79±0.01 | 0.72±0.01 | 0.50±0.01 | 0.28±0.01 | 0.25±0.00 | 0.21±0.00 | 0.29±0.01 | 0.68±0.03 | 1.63±0.05 | 0.25±0.01 | 0.51±0.01 | 0.38±0.01 | 0.26±0.01 | 0.74±0.01 | 2.05±0.04 | 2.00±0.03 | 0.36±0.01 | 0.23±0.01 |
[1.38, 1.63] | [0.78, 0.80] | [0.72, 0.73] | [0.50, 0.51] | [0.27, 0.29] | [0.24, 0.25] | [0.21, 0.21] | [0.28, 0.30] | [0.64, 0.71] | [1.58, 1.70] | [0.25, 0.26] | [0.50, 0.52] | [0.37, 0.39] | [0.24, 0.28] | [0.73, 0.76] | [2.02, 2.10] | [1.98, 2.05] | [0.36, 0.37] | [0.22, 0.25] | ||
hova | Minor (n= 27) | 1.55±0.10 | 0.66±0.03 | 0.66±0.02 | 0.49±0.02 | 0.25±0.02 | 0.27±0.01 | 0.23±0.01 | 0.29±0.01 | 0.73±0.03 | 1.53±0.08 | 0.30±0.01 | 0.50±0.01 | 0.37±0.02 | 0.27±0.02 | 0.75±0.02 | 1.68±0.09 | 1.92±0.06 | 0.38±0.02 | 0.25±0.02 |
[1.40, 1.74] | [0.63, 0.71] | [0.63, 0.70] | [0.47, 0.52] | [0.22, 0.28] | [0.24, 0.30] | [0.21, 0.25] | [0.27, 0.30] | [0.67, 0.78] | [1.41, 1.83] | [0.28, 0.32] | [0.48, 0.52] | [0.34, 0.41] | [0.23, 0.30] | [0.71, 0.78] | [1.51, 1.86] | [1.80, 2.04] | [0.34, 0.42] | [0.19, 0.29] | ||
Major (n= 10) | 3.17±0.16 | 0.94±0.03 | 0.77±0.01 | 0.53±0.01 | 0.27±0.01 | 0.26±0.01 | 0.22±0.01 | 0.30±0.01 | 0.86±0.02 | 0.87±0.04 | 0.21±0.01 | 0.47±0.01 | 0.58±0.02 | 0.22±0.01 | 0.55±0.02 | 1.06±0.06 | 0.97±0.04 | 1.34±0.04 | 0.18±0.01 | |
[2.94, 3.48] | [0.89, 1.01] | [0.75, 0.79] | [0.52, 0.55] | [0.26, 0.28] | [0.23, 0.27] | [0.21, 0.23] | [0.29, 0.31] | [0.83, 0.90] | [0.80, 0.92] | [0.19, 0.23] | [0.46, 0.48] | [0.55, 0.62] | [0.21, 0.24] | [0.54, 0.58] | [0.95, 1.12] | [0.91, 1.02] | [1.30, 1.46] | [0.17, 0.19] | ||
hovahovoides | Minor (n= 55) | 1.56±0.13 | 0.65±0.02 | 0.67±0.02 | 0.47±0.01 | 0.28±0.01 | 0.30±0.01 | 0.25±0.01 | 0.29±0.01 | 0.75±0.03 | 1.54±0.08 | 0.30±0.01 | 0.49±0.01 | 0.35±0.01 | 0.29±0.02 | 0.72±0.02 | 1.77±0.09 | 1.91±0.06 | 0.35±0.01 | 0.30±0.02 |
[1.16, 1.83] | [0.59, 0.68] | [0.63, 0.71] | [0.43, 0.49] | [0.25, 0.31] | [0.28, 0.33] | [0.22, 0.26] | [0.27, 0.31] | [0.69, 0.83] | [1.43, 1.75] | [0.27, 0.34] | [0.46, 0.51] | [0.32, 0.38] | [0.25, 0.34] | [0.67, 0.76] | [1.62, 2.00] | [1.79, 2.11] | [0.33, 0.39] | [0.25, 0.35] | ||
Major (n= 17) | 2.66±0.20 | 0.90±0.04 | 0.77±0.02 | 0.50±0.01 | 0.29±0.01 | 0.29±0.02 | 0.23±0.01 | 0.32±0.01 | 0.93±0.04 | 0.92±0.07 | 0.22±0.01 | 0.45±0.02 | 0.56±0.02 | 0.22±0.01 | 0.52±0.02 | 1.14±0.06 | 1.00±0.04 | 1.37±0.03 | 0.21±0.01 | |
[2.40, 3.07] | [0.85, 0.98] | [0.72, 0.80] | [0.48, 0.51] | [0.27, 0.32] | [0.25, 0.31] | [0.20, 0.24] | [0.30, 0.33] | [0.88, 1.01] | [0.75, 1.01] | [0.20, 0.23] | [0.43, 0.48] | [0.54, 0.60] | [0.20, 0.26] | [0.47, 0.55] | [1.01, 1.21] | [0.91, 1.06] | [1.32, 1.42] | [0.18, 0.23] | ||
imitator | Minor (n= 44) | 2.42±0.88 | 0.85±0.10 | 0.76±0.04 | 0.54±0.02 | 0.27±0.02 | 0.26±0.01 | 0.23±0.01 | 0.28±0.02 | 0.71±0.07 | 1.21±0.36 | 0.21±0.03 | 0.45±0.04 | 0.35±0.06 | 0.23±0.02 | 0.62±0.05 | 1.69±0.46 | 1.67±0.34 | 0.28±0.04 | 0.22±0.03 |
[1.34, 3.94] | [0.72, 1.03] | [0.68, 0.84] | [0.42, 0.57] | [0.22, 0.32] | [0.21, 0.28] | [0.16, 0.25] | [0.23, 0.32] | [0.53, 0.83] | [0.71, 1.73] | [0.16, 0.27] | [0.33, 0.51] | [0.27, 0.46] | [0.16, 0.26] | [0.49, 0.70] | [1.01, 2.39] | [1.17, 2.15] | [0.22, 0.35] | [0.16, 0.27] | ||
immaculatus | Minor (n= 12) | 1.20±0.13 | 0.79±0.03 | 0.78±0.02 | 0.51±0.01 | 0.26±0.02 | 0.32±0.01 | 0.26±0.01 | 0.30±0.01 | 0.70±0.03 | 1.28±0.05 | 0.25±0.01 | 0.48±0.01 | 0.44±0.01 | 0.27±0.01 | 0.74±0.01 | 1.34±0.05 | 1.66±0.06 | 0.35±0.01 | 0.21±0.02 |
[1.01, 1.45] | [0.73, 0.83] | [0.73, 0.80] | [0.48, 0.53] | [0.22, 0.28] | [0.30, 0.33] | [0.24, 0.28] | [0.29, 0.32] | [0.65, 0.74] | [1.19, 1.34] | [0.23, 0.28] | [0.47, 0.50] | [0.41, 0.47] | [0.24, 0.29] | [0.72, 0.76] | [1.26, 1.41] | [1.56, 1.73] | [0.33, 0.37] | [0.18, 0.24] | ||
Major (n= 7) | 2.32±0.12 | 1.00±0.03 | 0.85±0.01 | 0.53±0.01 | 0.28±0.02 | 0.32±0.01 | 0.25±0.01 | 0.32±0.01 | 0.84±0.01 | 0.82±0.03 | 0.19±0.00 | 0.47±0.01 | 0.65±0.01 | 0.23±0.01 | 0.52±0.02 | 0.95±0.05 | 0.95±0.01 | 1.33±0.02 | 0.16±0.01 | |
[2.13, 2.46] | [0.96, 1.04] | [0.83, 0.86] | [0.52, 0.56] | [0.25, 0.30] | [0.31, 0.33] | [0.24, 0.26] | [0.31, 0.33] | [0.82, 0.86] | [0.76, 0.87] | [0.19, 0.19] | [0.45, 0.48] | [0.63, 0.66] | [0.22, 0.25] | [0.50, 0.55] | [0.85, 1.00] | [0.93, 0.97] | [1.30, 1.36] | [0.14, 0.17] | ||
joany | Minor (n= 2) | 1.54±0.09 | 0.73±0.01 | 0.68±0.01 | 0.51±0.00 | 0.26±0.00 | 0.27±0.00 | 0.22±0.00 | 0.28±0.00 | 0.68±0.02 | 1.62±0.01 | 0.27±0.00 | 0.50±0.00 | 0.39±0.02 | 0.28±0.00 | 0.76±0.00 | 1.95±0.01 | 2.00±0.00 | 0.39±0.00 | 0.23±0.01 |
[1.48, 1.60] | [0.72, 0.73] | [0.68, 0.69] | [0.50, 0.51] | [0.26, 0.26] | [0.27, 0.27] | [0.22, 0.23] | [0.28, 0.29] | [0.67, 0.69] | [1.61, 1.63] | [0.27, 0.28] | [0.49, 0.50] | [0.38, 0.40] | [0.28, 0.28] | [0.76, 0.77] | [1.95, 1.96] | [2.00, 2.01] | [0.39, 0.40] | [0.22, 0.23] | ||
karsti | Minor (n= 2) | 1.83±0.09 | 0.70±0.00 | 0.73±0.01 | 0.52±0.00 | 0.26±0.01 | 0.25±0.00 | 0.21±0.00 | 0.30±0.00 | 0.75±0.01 | 1.65±0.03 | 0.26±0.01 | 0.53±0.00 | 0.37±0.01 | 0.21±0.00 | 0.42±0.01 | 2.08±0.02 | 1.93±0.02 | 0.26±0.00 | 0.24±0.00 |
[1.77, 1.89] | [0.70, 0.70] | [0.72, 0.73] | [0.52, 0.52] | [0.25, 0.27] | [0.25, 0.25] | [0.21, 0.22] | [0.29, 0.30] | [0.75, 0.76] | [1.63, 1.67] | [0.25, 0.26] | [0.53, 0.53] | [0.36, 0.37] | [0.20, 0.21] | [0.41, 0.42] | [2.06, 2.09] | [1.92, 1.94] | [0.26, 0.26] | [0.24, 0.24] | ||
kelimaso | Minor (n= 17) | 1.68±0.12 | 0.84±0.02 | 0.79±0.02 | 0.54±0.06 | 0.28±0.01 | 0.31±0.02 | 0.26±0.01 | 0.35±0.01 | 0.81±0.03 | 1.19±0.05 | 0.19±0.01 | 0.49±0.01 | 0.42±0.01 | 0.26±0.02 | 0.72±0.02 | 1.27±0.04 | 1.55±0.06 | 0.38±0.01 | 0.25±0.01 |
[1.51, 1.88] | [0.80, 0.88] | [0.76, 0.83] | [0.31, 0.57] | [0.27, 0.29] | [0.28, 0.34] | [0.25, 0.27] | [0.33, 0.37] | [0.75, 0.88] | [1.11, 1.27] | [0.16, 0.21] | [0.46, 0.51] | [0.38, 0.44] | [0.22, 0.28] | [0.65, 0.74] | [1.20, 1.38] | [1.45, 1.73] | [0.35, 0.40] | [0.22, 0.27] | ||
Major (n= 7) | 2.83±0.25 | 0.99±0.04 | 0.85±0.01 | 0.54±0.01 | 0.31±0.01 | 0.31±0.01 | 0.25±0.01 | 0.35±0.01 | 0.92±0.04 | 0.82±0.07 | 0.15±0.01 | 0.45±0.01 | 0.67±0.02 | 0.23±0.02 | 0.56±0.02 | 0.93±0.06 | 0.89±0.06 | 1.35±0.03 | 0.20±0.02 | |
[2.35, 3.15] | [0.92, 1.04] | [0.84, 0.87] | [0.53, 0.56] | [0.30, 0.32] | [0.30, 0.33] | [0.24, 0.27] | [0.33, 0.37] | [0.90, 1.01] | [0.77, 0.96] | [0.14, 0.17] | [0.44, 0.46] | [0.65, 0.70] | [0.21, 0.25] | [0.52, 0.60] | [0.88, 1.06] | [0.84, 1.02] | [1.30, 1.39] | [0.16, 0.24] | ||
liandia | Minor (n= 44) | 1.04±0.11 | 0.77±0.02 | 0.76±0.02 | 0.51±0.01 | 0.26±0.02 | 0.30±0.01 | 0.24±0.01 | 0.29±0.01 | 0.66±0.03 | 1.22±0.08 | 0.25±0.02 | 0.48±0.01 | 0.44±0.02 | 0.26±0.01 | 0.73±0.02 | 1.27±0.07 | 1.64±0.07 | 0.35±0.02 | 0.21±0.02 |
[0.88, 1.53] | [0.73, 0.85] | [0.69, 0.79] | [0.48, 0.54] | [0.23, 0.29] | [0.27, 0.34] | [0.22, 0.27] | [0.27, 0.31] | [0.59, 0.76] | [0.87, 1.33] | [0.20, 0.29] | [0.46, 0.51] | [0.37, 0.48] | [0.23, 0.30] | [0.68, 0.75] | [0.98, 1.36] | [1.38, 1.78] | [0.32, 0.40] | [0.16, 0.24] | ||
Major (n= 12) | 1.95±0.16 | 0.89±0.01 | 0.78±0.01 | 0.53±0.01 | 0.29±0.01 | 0.30±0.01 | 0.23±0.01 | 0.30±0.00 | 0.84±0.03 | 0.75±0.06 | 0.19±0.01 | 0.47±0.01 | 0.66±0.01 | 0.24±0.01 | 0.53±0.01 | 0.92±0.05 | 0.94±0.01 | 1.34±0.02 | 0.18±0.01 | |
[1.73, 2.15] | [0.85, 0.93] | [0.73, 0.84] | [0.52, 0.55] | [0.27, 0.32] | [0.28, 0.31] | [0.21, 0.25] | [0.29, 0.32] | [0.76, 0.91] | [0.69, 0.86] | [0.17, 0.19] | [0.45, 0.48] | [0.64, 0.70] | [0.21, 0.27] | [0.48, 0.58] | [0.83, 0.99] | [0.92, 0.97] | [1.28, 1.37] | [0.16, 0.19] | ||
lokobe | Minor (n= 8) | 1.65±0.05 | 0.64±0.01 | 0.64±0.01 | 0.46±0.01 | 0.29±0.01 | 0.26±0.01 | 0.22±0.01 | 0.26±0.01 | 0.72±0.02 | 1.80±0.03 | 0.30±0.01 | 0.50±0.01 | 0.29±0.00 | 0.21±0.00 | 0.67±0.01 | 2.28±0.04 | 2.11±0.02 | 0.29±0.01 | 0.26±0.01 |
[1.60, 1.72] | [0.62, 0.66] | [0.63, 0.66] | [0.44, 0.48] | [0.28, 0.31] | [0.25, 0.27] | [0.22, 0.23] | [0.25, 0.27] | [0.69, 0.75] | [1.76, 1.85] | [0.27, 0.31] | [0.49, 0.51] | [0.28, 0.29] | [0.21, 0.22] | [0.65, 0.68] | [2.24, 2.33] | [2.09, 2.14] | [0.29, 0.30] | [0.24, 0.27] | ||
mahafaly | Minor (n= 19) | 1.05±0.05 | 0.70±0.02 | 0.74±0.02 | 0.50±0.01 | 0.23±0.01 | 0.27±0.01 | 0.23±0.01 | 0.27±0.01 | 0.62±0.02 | 1.40±0.04 | 0.32±0.01 | 0.44±0.01 | 0.38±0.03 | 0.25±0.01 | 0.72±0.06 | 1.56±0.02 | 1.81±0.12 | 0.37±0.03 | 0.25±0.02 |
[0.93, 1.13] | [0.65, 0.73] | [0.71, 0.78] | [0.48, 0.52] | [0.20, 0.24] | [0.25, 0.28] | [0.20, 0.24] | [0.26, 0.28] | [0.58, 0.65] | [1.33, 1.45] | [0.30, 0.35] | [0.43, 0.47] | [0.35, 0.50] | [0.23, 0.28] | [0.67, 0.97] | [1.52, 1.60] | [1.35, 1.90] | [0.34, 0.50] | [0.23, 0.29] | ||
mixtellus | Minor (n= 25) | 1.59±0.10 | 0.63±0.02 | 0.64±0.02 | 0.46±0.01 | 0.30±0.01 | 0.28±0.01 | 0.23±0.01 | 0.28±0.01 | 0.76±0.04 | 1.55±0.07 | 0.30±0.02 | 0.49±0.01 | 0.37±0.01 | 0.29±0.01 | 0.75±0.02 | 1.77±0.08 | 1.95±0.05 | 0.39±0.02 | 0.28±0.02 |
[1.36, 1.75] | [0.58, 0.66] | [0.62, 0.68] | [0.44, 0.49] | [0.28, 0.31] | [0.26, 0.30] | [0.21, 0.24] | [0.26, 0.31] | [0.69, 0.85] | [1.40, 1.73] | [0.27, 0.33] | [0.46, 0.51] | [0.34, 0.39] | [0.27, 0.30] | [0.71, 0.78] | [1.64, 1.92] | [1.85, 2.06] | [0.36, 0.42] | [0.23, 0.30] | ||
Major (n= 8) | 2.92±0.15 | 0.93±0.03 | 0.77±0.02 | 0.50±0.01 | 0.29±0.02 | 0.28±0.01 | 0.22±0.01 | 0.30±0.01 | 0.93±0.02 | 0.82±0.05 | 0.22±0.01 | 0.44±0.01 | 0.58±0.01 | 0.22±0.01 | 0.54±0.01 | 1.05±0.05 | 0.97±0.02 | 1.32±0.03 | 0.19±0.01 | |
[2.62, 3.10] | [0.90, 0.99] | [0.75, 0.80] | [0.48, 0.52] | [0.27, 0.31] | [0.26, 0.29] | [0.20, 0.23] | [0.28, 0.31] | [0.89, 0.96] | [0.77, 0.91] | [0.21, 0.24] | [0.43, 0.48] | [0.56, 0.60] | [0.20, 0.23] | [0.53, 0.56] | [0.98, 1.12] | [0.95, 1.01] | [1.28, 1.37] | [0.17, 0.20] | ||
niavo | Minor (n= 8) | 2.20±0.15 | 0.59±0.03 | 0.61±0.03 | 0.50±0.03 | 0.28±0.02 | 0.25±0.01 | 0.22±0.01 | 0.30±0.02 | 0.81±0.04 | 1.70±0.11 | 0.27±0.01 | 0.51±0.04 | 0.34±0.03 | 0.29±0.02 | 0.72±0.01 | 2.05±0.17 | 2.05±0.12 | 0.37±0.01 | 0.30±0.02 |
[1.98, 2.38] | [0.55, 0.61] | [0.57, 0.65] | [0.46, 0.53] | [0.25, 0.32] | [0.24, 0.28] | [0.21, 0.23] | [0.28, 0.34] | [0.74, 0.87] | [1.59, 1.88] | [0.26, 0.30] | [0.44, 0.56] | [0.28, 0.37] | [0.26, 0.32] | [0.71, 0.74] | [1.87, 2.34] | [1.93, 2.26] | [0.35, 0.38] | [0.28, 0.32] | ||
Major (n= 7) | 4.02±0.35 | 0.92±0.08 | 0.73±0.04 | 0.53±0.01 | 0.28±0.01 | 0.24±0.01 | 0.20±0.01 | 0.31±0.01 | 0.90±0.05 | 0.95±0.12 | 0.19±0.01 | 0.48±0.02 | 0.55±0.02 | 0.22±0.02 | 0.54±0.01 | 1.25±0.16 | 1.00±0.08 | 1.34±0.04 | 0.21±0.02 | |
[3.69, 4.61] | [0.79, 1.01] | [0.66, 0.76] | [0.52, 0.55] | [0.26, 0.30] | [0.24, 0.26] | [0.20, 0.21] | [0.29, 0.33] | [0.83, 0.97] | [0.81, 1.19] | [0.18, 0.22] | [0.46, 0.51] | [0.52, 0.58] | [0.20, 0.26] | [0.53, 0.55] | [1.11, 1.55] | [0.93, 1.17] | [1.31, 1.41] | [0.18, 0.25] | ||
quadrimaculatus | Minor (n= 72) | 1.18±0.14 | 0.84±0.03 | 0.82±0.02 | 0.51±0.02 | 0.25±0.01 | 0.34±0.01 | 0.28±0.01 | 0.30±0.01 | 0.66±0.04 | 1.23±0.07 | 0.26±0.02 | 0.48±0.01 | 0.47±0.01 | 0.28±0.01 | 0.75±0.02 | 1.22±0.07 | 1.58±0.07 | 0.37±0.02 | 0.21±0.02 |
[0.86, 1.58] | [0.79, 0.93] | [0.79, 0.86] | [0.48, 0.54] | [0.22, 0.28] | [0.31, 0.37] | [0.25, 0.30] | [0.27, 0.35] | [0.58, 0.74] | [1.01, 1.36] | [0.21, 0.30] | [0.44, 0.51] | [0.44, 0.51] | [0.25, 0.31] | [0.62, 0.78] | [1.04, 1.38] | [1.40, 1.70] | [0.33, 0.41] | [0.16, 0.25] | ||
Major (n= 8) | 2.44±0.14 | 1.04±0.03 | 0.88±0.02 | 0.53±0.01 | 0.28±0.02 | 0.34±0.00 | 0.26±0.00 | 0.32±0.01 | 0.84±0.04 | 0.78±0.03 | 0.20±0.01 | 0.46±0.01 | 0.69±0.02 | 0.24±0.02 | 0.54±0.03 | 0.88±0.02 | 0.87±0.04 | 1.26±0.14 | 0.16±0.01 | |
[2.19, 2.66] | [1.00, 1.08] | [0.85, 0.90] | [0.50, 0.54] | [0.26, 0.31] | [0.34, 0.35] | [0.26, 0.27] | [0.30, 0.33] | [0.77, 0.88] | [0.75, 0.82] | [0.19, 0.21] | [0.44, 0.47] | [0.65, 0.72] | [0.21, 0.28] | [0.51, 0.58] | [0.85, 0.91] | [0.81, 0.93] | [0.95, 1.36] | [0.15, 0.18] | ||
radamae | Minor (n= 24) | 1.27±0.09 | 0.65±0.02 | 0.69±0.02 | 0.46±0.01 | 0.29±0.01 | 0.32±0.01 | 0.26±0.01 | 0.29±0.01 | 0.71±0.03 | 1.52±0.07 | 0.31±0.01 | 0.49±0.01 | 0.34±0.02 | 0.25±0.01 | 0.70±0.01 | 1.69±0.07 | 1.87±0.06 | 0.34±0.01 | 0.28±0.01 |
[1.04, 1.45] | [0.62, 0.69] | [0.67, 0.72] | [0.44, 0.48] | [0.27, 0.31] | [0.30, 0.34] | [0.26, 0.27] | [0.28, 0.31] | [0.66, 0.75] | [1.42, 1.64] | [0.29, 0.34] | [0.46, 0.51] | [0.30, 0.37] | [0.23, 0.29] | [0.67, 0.72] | [1.59, 1.80] | [1.78, 1.99] | [0.31, 0.36] | [0.26, 0.30] | ||
Major (n= 7) | 2.68±0.31 | 0.95±0.04 | 0.80±0.02 | 0.50±0.01 | 0.30±0.01 | 0.30±0.01 | 0.24±0.01 | 0.30±0.01 | 0.88±0.03 | 0.80±0.05 | 0.21±0.01 | 0.44±0.01 | 0.59±0.01 | 0.21±0.01 | 0.54±0.02 | 1.00±0.04 | 0.88±0.02 | 1.37±0.02 | 0.19±0.02 | |
[2.29, 3.02] | [0.89, 0.99] | [0.77, 0.85] | [0.49, 0.51] | [0.29, 0.32] | [0.27, 0.31] | [0.22, 0.25] | [0.30, 0.32] | [0.82, 0.91] | [0.75, 0.89] | [0.20, 0.22] | [0.42, 0.45] | [0.56, 0.60] | [0.20, 0.22] | [0.52, 0.57] | [0.94, 1.04] | [0.86, 0.92] | [1.34, 1.41] | [0.17, 0.22] | ||
roeseli | Minor (n= 29) | 2.12±0.16 | 0.64±0.02 | 0.60±0.01 | 0.50±0.01 | 0.28±0.01 | 0.24±0.01 | 0.22±0.01 | 0.31±0.01 | 0.77±0.03 | 1.70±0.08 | 0.27±0.01 | 0.52±0.01 | 0.35±0.02 | 0.28±0.01 | 0.75±0.02 | 1.92±0.08 | 2.01±0.06 | 0.39±0.02 | 0.25±0.02 |
[1.82, 2.53] | [0.62, 0.69] | [0.57, 0.63] | [0.47, 0.52] | [0.26, 0.29] | [0.23, 0.27] | [0.21, 0.24] | [0.29, 0.33] | [0.73, 0.84] | [1.50, 1.81] | [0.25, 0.29] | [0.50, 0.54] | [0.32, 0.39] | [0.26, 0.31] | [0.71, 0.79] | [1.77, 2.05] | [1.88, 2.11] | [0.35, 0.43] | [0.20, 0.29] | ||
Major (n= 10) | 4.07±0.19 | 0.89±0.04 | 0.69±0.02 | 0.55±0.01 | 0.27±0.01 | 0.23±0.01 | 0.20±0.00 | 0.32±0.01 | 0.88±0.02 | 0.98±0.05 | 0.19±0.01 | 0.48±0.01 | 0.56±0.02 | 0.21±0.01 | 0.54±0.04 | 1.19±0.08 | 0.99±0.04 | 1.35±0.03 | 0.18±0.01 | |
[3.81, 4.40] | [0.85, 0.96] | [0.66, 0.73] | [0.53, 0.56] | [0.26, 0.29] | [0.22, 0.24] | [0.20, 0.21] | [0.31, 0.33] | [0.85, 0.91] | [0.91, 1.04] | [0.18, 0.20] | [0.45, 0.50] | [0.53, 0.60] | [0.20, 0.23] | [0.44, 0.56] | [1.07, 1.27] | [0.92, 1.05] | [1.30, 1.40] | [0.16, 0.20] | ||
rotrae | Minor (n= 18) | 1.29±0.09 | 0.85±0.05 | 0.80±0.06 | 0.53±0.01 | 0.25±0.01 | 0.31±0.01 | 0.25±0.01 | 0.31±0.01 | 0.66±0.03 | 1.19±0.05 | 0.24±0.01 | 0.47±0.02 | 0.44±0.01 | 0.26±0.01 | 0.74±0.01 | 1.25±0.04 | 1.60±0.07 | 0.38±0.01 | 0.20±0.01 |
[1.08, 1.42] | [0.81, 1.05] | [0.77, 1.05] | [0.51, 0.55] | [0.23, 0.25] | [0.29, 0.33] | [0.23, 0.26] | [0.29, 0.33] | [0.60, 0.71] | [1.08, 1.27] | [0.22, 0.26] | [0.40, 0.50] | [0.43, 0.47] | [0.23, 0.28] | [0.72, 0.75] | [1.19, 1.31] | [1.43, 1.76] | [0.36, 0.41] | [0.18, 0.24] | ||
Major (n= 4) | 2.29±0.17 | 1.03±0.01 | 0.84±0.03 | 0.56±0.01 | 0.27±0.02 | 0.31±0.01 | 0.24±0.01 | 0.33±0.01 | 0.79±0.02 | 0.75±0.03 | 0.18±0.01 | 0.46±0.01 | 0.67±0.01 | 0.22±0.01 | 0.55±0.01 | 0.88±0.03 | 0.88±0.04 | 1.37±0.03 | 0.14±0.01 | |
[2.03, 2.42] | [1.02, 1.04] | [0.82, 0.88] | [0.54, 0.57] | [0.24, 0.28] | [0.30, 0.32] | [0.24, 0.25] | [0.32, 0.33] | [0.77, 0.81] | [0.71, 0.79] | [0.17, 0.19] | [0.45, 0.47] | [0.66, 0.67] | [0.21, 0.23] | [0.54, 0.56] | [0.85, 0.91] | [0.85, 0.94] | [1.34, 1.41] | [0.13, 0.16] | ||
sambiranoensis | Minor (n= 8) | 1.77±0.11 | 0.60±0.01 | 0.58±0.01 | 0.47±0.01 | 0.32±0.04 | 0.22±0.01 | 0.18±0.01 | 0.28±0.01 | 0.75±0.03 | 2.04±0.07 | 0.28±0.01 | 0.51±0.01 | 0.30±0.02 | 0.23±0.01 | 0.70±0.02 | 2.36±0.06 | 2.22±0.04 | 0.33±0.03 | 0.27±0.02 |
[1.56, 1.89] | [0.59, 0.62] | [0.56, 0.60] | [0.46, 0.49] | [0.30, 0.42] | [0.21, 0.23] | [0.17, 0.19] | [0.26, 0.30] | [0.70, 0.79] | [1.97, 2.18] | [0.27, 0.29] | [0.49, 0.52] | [0.27, 0.32] | [0.21, 0.26] | [0.67, 0.72] | [2.29, 2.46] | [2.18, 2.30] | [0.28, 0.36] | [0.25, 0.31] | ||
Major (n= 1) | 4.45 | 0.98 | 0.74 | 0.52 | 0.30 | 0.25 | 0.21 | 0.31 | 0.88 | 0.84 | 0.18 | 0.46 | 0.57 | 0.22 | 0.53 | 1.12 | 0.93 | 1.29 | 0.20 | |
strangulatus | Minor (n= 18) | 1.59±0.11 | 0.67±0.02 | 0.66±0.01 | 0.50±0.01 | 0.27±0.01 | 0.25±0.01 | 0.22±0.01 | 0.30±0.01 | 0.80±0.03 | 1.58±0.05 | 0.28±0.01 | 0.52±0.01 | 0.33±0.01 | 0.24±0.02 | 0.68±0.01 | 1.87±0.05 | 1.94±0.03 | 0.33±0.01 | 0.25±0.02 |
[1.41, 1.83] | [0.63, 0.69] | [0.64, 0.69] | [0.48, 0.52] | [0.26, 0.29] | [0.24, 0.27] | [0.21, 0.23] | [0.26, 0.31] | [0.71, 0.85] | [1.50, 1.66] | [0.26, 0.30] | [0.51, 0.56] | [0.31, 0.35] | [0.22, 0.27] | [0.65, 0.69] | [1.80, 2.01] | [1.86, 1.99] | [0.31, 0.36] | [0.19, 0.27] | ||
Major (n= 6) | 3.22±0.36 | 0.91±0.06 | 0.75±0.02 | 0.54±0.02 | 0.32±0.11 | 0.25±0.01 | 0.22±0.01 | 0.31±0.01 | 0.89±0.06 | 0.94±0.11 | 0.20±0.01 | 0.49±0.01 | 0.59±0.03 | 0.22±0.01 | 0.56±0.02 | 1.22±0.12 | 0.98±0.04 | 1.42±0.04 | 0.21±0.02 | |
[2.73, 3.74] | [0.80, 0.98] | [0.72, 0.78] | [0.51, 0.55] | [0.27, 0.54] | [0.24, 0.27] | [0.21, 0.23] | [0.30, 0.32] | [0.80, 0.93] | [0.82, 1.08] | [0.19, 0.23] | [0.48, 0.50] | [0.55, 0.64] | [0.21, 0.25] | [0.54, 0.59] | [1.06, 1.36] | [0.92, 1.03] | [1.38, 1.48] | [0.20, 0.23] | ||
tapia | Minor (n= 17) | 1.37±0.14 | 0.67±0.03 | 0.69±0.01 | 0.52±0.01 | 0.24±0.01 | 0.25±0.01 | 0.23±0.01 | 0.29±0.01 | 0.73±0.04 | 1.61±0.08 | 0.29±0.01 | 0.52±0.01 | 0.34±0.02 | 0.24±0.01 | 0.69±0.04 | 1.79±0.07 | 1.97±0.15 | 0.33±0.02 | 0.23±0.02 |
[1.17, 1.63] | [0.64, 0.72] | [0.68, 0.71] | [0.49, 0.54] | [0.23, 0.25] | [0.23, 0.27] | [0.21, 0.25] | [0.26, 0.31] | [0.67, 0.79] | [1.48, 1.71] | [0.28, 0.31] | [0.51, 0.53] | [0.25, 0.36] | [0.22, 0.26] | [0.54, 0.72] | [1.67, 1.91] | [1.83, 2.52] | [0.25, 0.36] | [0.19, 0.26] | ||
Major (n= 5) | 3.20±0.21 | 0.99±0.03 | 0.81±0.01 | 0.52±0.01 | 0.29±0.02 | 0.26±0.01 | 0.23±0.01 | 0.31±0.00 | 0.88±0.01 | 0.85±0.06 | 0.19±0.01 | 0.46±0.00 | 0.59±0.02 | 0.20±0.00 | 0.55±0.02 | 1.08±0.05 | 0.97±0.02 | 1.35±0.03 | 0.20±0.01 | |
[2.84, 3.38] | [0.94, 1.01] | [0.78, 0.82] | [0.50, 0.54] | [0.27, 0.32] | [0.25, 0.28] | [0.22, 0.24] | [0.31, 0.32] | [0.86, 0.89] | [0.81, 0.96] | [0.18, 0.21] | [0.46, 0.47] | [0.56, 0.61] | [0.20, 0.20] | [0.53, 0.59] | [1.04, 1.16] | [0.94, 1.00] | [1.31, 1.38] | [0.18, 0.21] | ||
tendryi | Minor (n= 2) | 1.57±0.01 | 0.54±0.04 | 0.55±0.03 | 0.44±0.00 | 0.31±0.02 | 0.25±0.01 | 0.20±0.00 | 0.26±0.02 | 0.77±0.02 | 1.95±0.00 | 0.28±0.01 | 0.51±0.00 | 0.28±0.01 | 0.25±0.02 | 0.69±0.01 | 2.28±0.01 | 2.08±0.06 | 0.30±0.00 | 0.28±0.01 |
[1.56, 1.58] | [0.52, 0.57] | [0.52, 0.57] | [0.44, 0.44] | [0.30, 0.33] | [0.24, 0.26] | [0.20, 0.21] | [0.24, 0.27] | [0.75, 0.78] | [1.95, 1.96] | [0.27, 0.29] | [0.51, 0.51] | [0.28, 0.29] | [0.24, 0.26] | [0.69, 0.70] | [2.28, 2.29] | [2.04, 2.12] | [0.30, 0.30] | [0.27, 0.29] | ||
vano | Minor (n= 2) | 1.03±0.01 | 0.56±0.01 | 0.62±0.00 | 0.43±0.01 | 0.34±0.00 | 0.29±0.00 | 0.25±0.01 | 0.27±0.00 | 0.74±0.00 | 1.68±0.03 | 0.29±0.00 | 0.47±0.01 | 0.28±0.00 | 0.24±0.02 | 0.68±0.01 | 1.92±0.06 | 1.98±0.09 | 0.29±0.02 | 0.25±0.01 |
[1.02, 1.03] | [0.55, 0.57] | [0.62, 0.62] | [0.42, 0.43] | [0.34, 0.34] | [0.29, 0.30] | [0.24, 0.25] | [0.26, 0.27] | [0.74, 0.74] | [1.66, 1.70] | [0.29, 0.29] | [0.46, 0.49] | [0.28, 0.29] | [0.22, 0.25] | [0.67, 0.69] | [1.88, 1.97] | [1.92, 2.04] | [0.28, 0.31] | [0.24, 0.25] | ||
Major (n= 1) | 1.67 | 0.64 | 0.62 | 0.46 | 0.34 | 0.28 | 0.25 | 0.30 | 0.92 | 1.00 | 0.24 | 0.50 | 0.45 | 0.21 | 0.46 | 1.32 | 1.11 | 1.42 | 0.20 |
In some species, the existing samples were below the threshold level to apply the quantitative, morphometry-based classification methods and were not included in the analysis. However, the measurement values of the individual specimens were used in the description of each of these species whose delimitations are supported by the qualitative morphological traits and other data from their biology and ecology.
Morphometric data of the major workers are also provided in the description but were excluded in statistical analysis.
As different allometric properties are present in the Malagasy Camponotus (see
Species hypotheses were generated using the Nest Centroid clustering (NC-clustering) technique. The procedure followed
To validate the species boundaries and reliability of the clusters recognized by the exploratory analyses, cumulative Linear Discriminant Analysis (LDA) was performed repeatedly until the final classification, showing the highest posterior probability values, was attained.
The NC-clustering dendrogram revealed 33 clusters, which are interpreted as 33 species (Fig.
Classification matrix of species showing the classification success (percentage), the observed classification (rows) and the predicted classification (columns). Numbers in the matrix are specimen counts.
Observed species | Predicted species | |||||||||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
aro | asara | atimo | aurosus | becki | bemaheva | boivini | bozaka | cemeryi | cervicalis | daraina | dufouri | gibber | gouldi | hagensii | hova | hovahovoides | imitator | immaculatus | kelimaso | liandia | lokobe | lubbocki | mahafaly | mixtellus | niavo | quadrimaculatus | radamae | roeseli | rotrae | sambiranoensis | strangulatus | tapia | Identification success (%) | |
aro | 10 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
asara | 0 | 18 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
atimo | 0 | 0 | 31 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
aurosus | 0 | 0 | 0 | 16 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
becki | 0 | 0 | 0 | 0 | 10 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 90.91 |
bemaheva | 0 | 0 | 0 | 0 | 0 | 22 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 95.65 |
boivini | 0 | 0 | 0 | 0 | 0 | 0 | 49 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
bozaka | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 11 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 91.67 |
cemeryi | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 16 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
cervicalis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 8 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
daraina | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 18 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 94.74 |
dufouri | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 19 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
gibber | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 33 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 0 | 0 | 0 | 0 | 0 | 0 | 91.67 |
gouldi | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 18 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
hagensii | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 16 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 94.12 |
hova | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 26 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 96.30 |
hovahovoides | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 55 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
imitator | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 43 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 97.73 |
immaculatus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 11 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 91.67 |
kelimaso | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
liandia | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 57 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
lokobe | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 8 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
lubbocki | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 16 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
mahafaly | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 19 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
mixtellus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 25 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
niavo | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 8 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 100 |
quadrimaculatus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 69 | 0 | 0 | 1 | 0 | 0 | 0 | 98.57 |
radamae | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 24 | 0 | 0 | 0 | 0 | 0 | 100 |
roeseli | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 29 | 0 | 0 | 0 | 0 | 100 |
rotrae | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 18 | 0 | 0 | 0 | 94.74 |
sambiranoensis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 8 | 0 | 0 | 100 |
strangulatus | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 0 | 94.44 |
tapia | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 100 |
Total | 10 | 18 | 32 | 16 | 10 | 23 | 49 | 11 | 16 | 8 | 18 | 19 | 34 | 19 | 16 | 27 | 55 | 43 | 11 | 17 | 58 | 8 | 16 | 20 | 25 | 8 | 73 | 24 | 29 | 19 | 8 | 17 | 19 | 98.20 |
Myrmosaga
Myrmopytia Emery, 1920a: 243 [as subgenus of Camponotus]. Type species: Camponotus imitator Forel, 1891: 209, by monotypy, by original designation. Syn. nov.
As for most Camponotus species, minor and major worker castes exist within a colony of the subgenus Myrmosaga. In addition, various worker forms showing continuous morphological variation between these two castes are observed. The following combination of characters can be used to reliably distinguish the two extreme worker castes in the subgenus Myrmosaga from other Malagasy subgenera and species groups of Camponotus.
Major worker similar to minor worker, but characterized by the following distinctive traits: larger, heart-shaped head; less protruding eye not breaking lateral cephalic margin; more robust mesosoma; stronger mandibles (armed with at least seven teeth and denticles); clypeus with anterolateral angle and straight anterior margin; antennal scape shorter, at most apical 1/3 extending beyond posterior margin of head; metanotum distinctly visible; petiolar node much higher than long (more flattened anteroposteriorly); more erect hairs on promesonotum and the junction of propodeal dorsum and declivity.
Originally
Species of the Malagasy Camponotus that have been previously placed in the subgenera Dinomyrmex and Myrmoturba by
Camponotus imitator and the following three species C. jodina, C. karaha, and C. longicollis have morphologically similar minor worker caste and have been grouped together in the subgenus Myrmopytia (
aina sp. nov.
aro sp. nov.
asara sp. nov.
atimo sp. nov.
aurosus Roger, 1863
becki Santschi, 1923, stat. nov.
bemaheva sp. nov.
boivini Forel, 1891, stat. rev.
= maculatus st. fairmairei Santschi, 1911a, syn. nov.
bozaka sp. nov.
cemeryi Özdikmen, 2010, stat. nov.
cervicalis Roger, 1863
= gaullei Santschi, 1911a, syn. nov.
= perroti Forel, 1897, syn. nov.
= perroti aeschylus Forel, 1913, syn. nov.
= gerberti Donisthorpe, 1949, syn. nov.
daraina Rakotonirina & Fisher, sp. nov.
dufouri Forel, 1891
= dufouri imerinensis Forel, 1891, syn. nov.
gibber Forel, 1891
gouldi Forel, 1886a
hagensii Forel, 1886a
harenarum sp. nov.
hova Forel, 1891
= hova var. obscuratus Emery, 1925, syn. nov.
hovahovoides Forel, 1892
= radamae var. hovoides Dalla Torre, 1893, syn. nov.
imitator Forel, 1891
immaculatus Forel, 1892
= quadrimaculatus opacata Emery, 1925, syn. nov.
joany sp. nov.
karsti sp. nov.
kelimaso sp. nov.
liandia Rakotonirina & Fisher, 2018
lokobe sp. nov.
lubbocki Forel, 1886b
mahafaly sp. nov.
mixtellus Forel, 1891, stat. nov.
niavo sp. nov.
quadrimaculatus Forel, 1886a
= kelleri Forel, 1886b, syn. nov.
= kelleri var. invalidus Forel, 1897, syn. nov.
= quadrimaculatus sellaris Emery, 1895, syn. nov.
radamae Forel, 1891, stat. rev.
roeseli Forel, 1910
= maculatus st. legionarium Santschi, 1911b, syn. nov.
rotrae sp. nov.
sambiranoensis sp. nov.
strangulatus Santschi, 1911a
= hova maculatoides Emery, 1920b, syn. nov.
tapia sp. nov.
tendryi sp. nov.
vano sp. nov.
Most of the species names listed above correspond to different species codes that have been previously used on AntWeb and in publications. These changes are provided in the present study (Table
List of the species names and their previous corresponding species codes used on AntWeb and in publications.
Species name | Species code |
---|---|
aina | MG107 |
aro | MG072 |
asara | MG070 |
atimo | MG062 |
becki | MG071 |
bemaheva | MG064B |
boivini | MG045, MG049B, MG050 |
bozaka | MG060 |
cemeryi | MG046 |
cervicalis | MG065 |
daraina | MG064 |
dufouri | MG067, MG069 |
gibber | MG014, MG030 |
harenarum | MG105 |
hova | MG056, MG056A, europa_sp1 |
hovohovoides | MG054, MG055 |
immaculatus | MG013 |
joany | MG108 |
karsti | MG106 |
kelimaso | MG056B |
liandia | MG010, MG012 |
lokobe | MG068 |
mahafaly | MG062B |
mixtellus | MG053 |
niavo | MG057 |
quadrimaculatus | MG016 |
radamae | MG048 |
roeseli | MG059 |
rotrae | MG015 |
sambiranoensis | MG066 |
tapia | MG061, MG063 |
tendryi | MG052 |
vano | MG125 |
1 | With head in full-face view, eyes not breaking lateral cephalic margins (Fig. |
2 |
– | With head in full-face view, eyes breaking lateral cephalic margins (Fig. |
5 |
2 | Dorsal margin of propodeum entirely straight (Fig. |
3 |
– | Dorsal margin of propodeum with blunt angle at ca. posterior 1/2 (Fig. |
4 |
3 | Propodeal dorsum 3 × as long as the declivity; petiolar node with dorsal margin as long as posterior margin (Fig. |
aina |
– | Propodeal dorsum 2 × as long as the declivity; petiolar node with dorsal margin shorter than posterior margin (Fig. |
joany |
4 | With mesosoma in lateral view, junction of mesonotum and anterior 1/2 of propodeum continuously straight, sloping down to make noticeable angle with posterior 1/2 of propodeum; petiolar node low and long (Fig. |
karsti |
– | With mesosoma in lateral view, mesonotum and anterior 1/2 of propodeum forming separate convexities; petiolar node ca. as high as long (Fig. |
harenarum |
5 | With head in full-face view, lateral cephalic margins converging posteriorly towards eye level (Fig. |
6 |
– | With head in full-face view, lateral cephalic margin either approximately parallel or diverging posteriorly towards eye level (Fig. |
14 |
6 | Two apical teeth of mandible normally spaced (Fig. |
7 |
– | Two apical teeth of mandible closely spaced (Fig. |
12 |
7 | With head in full-face view, anteromedian margin of clypeus broadly convex (Fig. |
8 |
– | With head in full-face view, anteromedian margin of clypeus either generally straight (Fig. |
10 |
8 | With head in full-face view, lateral cephalic margin anterior to eye level without erect hairs (Fig. |
sambiranoensis |
– | With head in full-face view, lateral cephalic margin anterior to eye level covered with erect hairs (Fig. |
9 |
9 | With head in full-face view, level of posterior ocular margin located approximately at posterior 1/4 of the length of head; antennal scape with appressed hairs (Fig. |
niavo |
– | With head in full-face view, level of posterior ocular margin located at posterior 1/3 of the length of head; antennal scape with suberect hairs (Fig. |
cervicalis |
10 | Anteromedian margin of clypeus noticeably excised medially (Fig. |
lokobe |
– | Anteromedian margin of clypeus straight (Fig. |
11 |
11 | Lateral cephalic margin posterior to eye level covered with erect hairs (Fig. |
dufouri |
– | Lateral cephalic margin posterior to eye level without erect hairs (Fig. |
tendryi |
12 | Lateral cephalic margin posterior to eye level without erect hairs (Fig. |
bemaheva |
– | Lateral cephalic margin posterior to eye level covered with erect hairs (Fig. |
13 |
13 | Integument of entire body yellowish orange to reddish orange (Fig. |
daraina |
– | Head and gaster reddish black and mesosoma reddish orange or integument entirely reddish black (Fig. |
roeseli |
14 | In full-face view, lateral margin of head anterior to eye level approximately parallel and covered with erect hairs (Fig. |
15 |
– | In full-face view, lateral margin of head anterior to eye level diverging posteriorly (Fig. |
23 |
15 | With mesosoma in lateral view, mesonotum elongate and constricted at midlength, propodeum broadly convex (Fig. |
imitator |
– | With mesosoma in lateral view, mesonotum short and lacking constriction at midlength; promesonotum an even convexity; propodeal dorsum almost straight (Fig. |
12 |
16 | Two apical teeth of mandible closely spaced (Fig. |
17 |
– | Two apical teeth of mandible normally spaced from each other (Fig. |
19 |
17 | Antennal scape covered with erect hairs; lateral cephalic margin posterior to eye level with erect hairs (Fig. |
hova |
– | Antennal scape covered with appressed hairs (Fig. |
18 |
18 | In lateral view, mesosoma much higher and short (MPH/ML: 0.34±0.01; 0.31–0.36), propodeal dorsum at most 3 × as long as declivity; petiolar node more or less flattened anteroposteriorly and tapering dorsally (Fig. |
radamae |
– | In lateral view, mesosoma very low and long (MPH/ML: 0.29±0.02; 0.28–0.31), propodeal dorsum at least four times as long as declivity; petiole much nodelike (Fig. |
vano |
19 | Antennal scape covered with appressed hairs (Fig. |
mahafaly |
– | Antennal scape covered with erect to suberect hairs (Fig. |
20 |
20 | Mesosoma generally short and high, its dorsal outline strongly convex, propodeal dorsum < 2 × height of declivity surface (Fig. |
cemeryi |
– | Mesosoma low and long, its dorsal outline more or less straight or not strongly convex, propodeal dorsum 2 × or more as long as height of declivity surface (Fig. |
21 |
21 | In lateral view, dorsum of mesosoma from mid-mesonotum to posterodorsal corner of propodeum approximately straight, propodeal dorsum ca. 3 × as long as height of declivity surface (Fig. |
hovahovoides |
– | In lateral view, posterior 1/2 of mesonotum to posterodorsal corner of propodeum slightly convex, propodeal dorsum ca. 2 × as long as height of declivity surface (Fig. |
22 |
22 | Antennal scape covered with suberect hairs inclined at ca. 30° (Fig. |
mixtellus |
– | Antennal scape covered with suberect hairs inclined at ca. 45° (Fig. |
boivini |
23 | With head in full-face view, lateral margins of head anterior to eye level parallel to each other (Fig. |
24 |
– | With head in full-face view, lateral margins of head anterior to eye level diverging posteriorly (Fig. |
31 |
24 | In full-face view, clypeus with distinctly visible anterolateral corner (Fig. |
25 |
– | In full-face view, clypeus without visible anterolateral corner, lateral and anteromedian clypeal margin continuously forming broad convexity (Fig. |
30 |
25 | Dark body color, or posterior portion of mesosoma pale brown; in lateral view, petiolar node scalelike or more or less compressed anteroposteriorly (Fig. |
26 |
– | Pale yellow to orange body color; in lateral view, petiole nodelike and not compressed anteroposteriorly (Fig. |
28 |
26 | With head in full-face view, anteromedian clypeal margin truncate; erect hairs present on lateral cephalic margin behind level of posterior ocular margin (Fig. |
becki |
– | With head in full-face view, anteromedian clypeal margin broadly convex or triangular; erect hairs absent on lateral cephalic margin behind level of posterior ocular margin (Fig. |
27 |
27 | In lateral view, mesosoma low and long, propodeal dorsum ca. 3 × as long as declivity, their junction rounded; petiolar node with dorsal margin inclined posteriorly; integument matte (Fig. |
asara |
– | In lateral view, mesosoma short and high, length of propodeal dorsum < 2 × height of declivity, their junction angulate; petiolar node scalelike; integument shining (Fig. |
bozaka |
28 | With head in oblique profile, three pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin | strangulatus |
– | With head in oblique profile, four or more pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin | 29 |
29 | With mesosoma in profile, junction of propodeal dorsum and declivity surface rounded (Fig. |
atimo |
– | With mesosoma in profile, junction of propodeal dorsum and declivity surface broadly angulate (Fig. |
tapia |
30 | In full-face view, posterior portion of head extending into broadly short neck (Fig. |
gouldi |
– | In full-face view, posterior portion of head normally rounded, not extending into a short neck (Fig. |
aro |
31 | With head in full-face view, lateral margin of head anterior to eye level with erect hairs (Fig. |
32 |
– | With head in full-face view, lateral margin of head anterior to eye level without erect hairs (Fig. |
33 |
32 | With head in full-face view, anterior clypeal margin broadly triangular (Fig. |
aurosus |
– | With head in full-face view, anterior clypeal margin truncate (Fig. |
hagensii |
33 | With head in full-face view, anterior clypeal margin broadly triangular (Fig. |
liandia |
– | With head in full-face view, anterior clypeal margin truncate (Fig. |
34 |
34 | No white spot on dorsum of second and third abdominal tergites (Fig. |
35 |
– | One pair of white spots on third abdominal tergites (Fig. |
37 |
35 | With mesosoma in lateral view, propodeal dorsum broadly concave (Fig. |
immaculatus |
– | With mesosoma in lateral view, propodeal dorsum approximately straight (Fig. |
36 |
36 | Eye small (EL/CL: 0.19±0.01; 0.16–0.21) (Fig. |
kelimaso |
– | Eye large (EL/CL: 0.24±0.01; 0.23–0.27) (Fig. |
lubbocki |
37 | Pronotum, mesonotum, and propodeum forming separate convexities, metanotal groove depressed; propodeum at lower level than promesonotum; petiolar node with dorsal face rounding to anterior and posterior faces (Fig. |
gibber |
– | Pronotum, mesonotum, and propodeum not forming separate convexities, metanotal groove not depressed; propodeum immediately in junction with promesonotum; petiolar node with dorsal face joining posterior face into an angle and rounding to anterior face (Fig. |
38 |
38 | Propodeal dorsum straight (Fig. |
rotrae |
– | Propodeal dorsum concave (Fig. |
quadrimaculatus |
Head in full-face view A C. joany (CASENT0408908) B C. radamae (CASENT0066777) C C. immaculatus (CASENT0179441).
Mesosoma in lateral view A C. aina (CASENT0217291) B C. harenarum (CASENT0499207).
Mesosoma and petiolar node in lateral view A C. aina (CASENT0217291) B C. joany (CASENT0408908).
Mesosoma and petiolar node in lateral view A C. karsti (CASENT0217292) B C. harenarum (CASENT0499207).
Head in full-face view A C. sambiranoensis (CASENT0231845) B C. radamae (CASENT0066777) C C. immaculatus (CASENT0179441).
Mandibles in frontal view A C. niavo (CASENT0300086) B C. bemaheva (CASENT0153778).
Head in full-face view A C. cervicalis (CASENT0217303) B C. dufouri (CASENT0487727) C C. lokobe (CASENT0436584).
Head in full-face view A C. sambiranoensis (CASENT0231845) B C. cervicalis (CASENT0217303).
Head in full-face view A C. niavo (CASENT0134004) B C. cervicalis (CASENT0217303).
Head in full-face view A C. lokobe (CASENT0436584) B C. tendryi (CASENT0145284).
Head in full-face view A C. dufouri (CASENT0274127) B C. tendryi (CASENT0145284).
Head in full-face view A C. bemaheva (CASENT0154158) B C. roeseli (CASENT0492473).
Body in lateral view A C. daraina (CASENT0077433) B C. roeseli (CASENT0492473).
Head in full-face view A C. mahafaly (CASENT0115287) B C. tapia (CASENT0493939) C C. quadrimaculatus (CASENT0096044).
Mesosoma in lateral view A C. imitator (CASENT0452849) B C. mahafaly (CASENT0115287).
Mandibles in frontal view A C. radamae (CASENT0228250) B C. boivini (CASENT0477264).
Head in full-face view A C. hova (CASENT0055710) B C. radamae (CASENT0217307).
Mesosoma and petiolar node in lateral view A C. radamae (CASENT066777) B C. vano (CASENT0217316).
Head in full-face view A C. mahafaly (CASENT0115287) B C. hovahovoides (CASENT0487242).
Mesosoma and petiolar node in lateral view A C. cemeryi (CASENT0217306) B C. hovahovoides (CASENT0496908).
Mesosoma in lateral view A C. hovahovoides (CASENT0496908) B C. mixtellus (CASENT0132606).
Head in full-face view A C. mixtellus (CASENT0076675) B C. boivini (CASENT0168345).
Head in full-face view A C. atimo (CASENT0454042) B C. quadrimaculatus (CASENT0096044).
Head in full-face view A C. tapia (CASENT0217310) B C. aro (CASENT0489917).
Posterior portion of mesosoma, petiolar node, and gaster in lateral view A C. bozaka (CASENT0217309) B C. atimo (CASENT0454042).
Head in full-face view A C. becki (CASENT0071380) B C. asara (CASENT0493252).
Mesosoma and petiolar node in lateral view A C. asara (CASENT0493252) B C. bozaka (CASENT0217309).
Head in lateral view A C. strangulatus (CASENT0136581) B C. tapia (CASENT0493939).
Mesosoma in lateral view A C. atimo (CASENT0454042) B C. tapia (CASENT0217310).
Head in full-face view A C. gouldi (CASENT0121617) B C. aro (CASENT0489917).
Head in full-face view A C. aurosus (CASENT0064815) B C. gibber (CASENT0188619).
Head in full-face view A C. aurosus (CASENT0064815) B C. hagensii (CASENT0191568).
Head in full-face view A C. liandia (CASENT0491145) B C. kelimaso (CASENT0487718).
Second and third abdominal tergites in dorsal view A C. kelimaso (CASENT0487718) B C. quadrimaculatus (CASENT0096044) C C. gibber (CASENT0491497).
Head, mesosoma and petiolar node in lateral view A C. immaculatus (CASENT0179441) B C. lubbocki (CASENT0486998).
Head in full-face view A C. kelimaso (CASENT0487718) B C. lubbocki (CASENT0486998).
Mesosoma and petiolar node in lateral view A C. gibber (CASENT0188619) B C. quadrimaculatus (CASENT0096044).
Mesosoma in lateral view A C. rotrae (CASENT0485625) B C. quadrimaculatus (CASENT0121629).
Madagascar: Province Antsiranana: RS Ankarana, 7 km SSE Matsaborimanga, -12.91667, 49.1, 150 m, tropical dry forest, on low vegetation, 28 Nov 1990 (P.S. Ward) collection code: PSW11019, specimen code: CASENT0217291 (
1 minor worker with same data as holotype but specimen coded as: CASENT0217290 (
Madagascar: Antsiranana: RS Ankarana, 7 km SSE Matsaborimanga, -12.91667, 49.1, 150 m, tropical dry forest (P.S. Ward) (
With head in full-face view, eye not breaking lateral cephalic margin; mesonotum short and lacking constriction; propodeal dorsum approximately straight, 3 × as long as the declivity; dorsal margin of petiole as long as posterior margin.
Minor worker. With head in full-face view, lateral margins anterior to eye level approximately parallel, rounding evenly towards slightly concave posterior margin; eye convex and large (EL/CS: 0.24±0.02; 0.22–0.26), not breaking lateral cephalic margin, location of its posterior margin at posterior 1/4 of head (PoOc/CL: 0.26±0.00; 0.26–0.26); frontal carinae parallel, widely opened posteriorly (FR/CS: 0.28±0.01; 0.27–0.29); clypeus with blunt or poorly defined anterolateral angle, anteromedian margin broadly convex; two apical teeth of mandible normally spaced; antennal scape relatively long (SL/CS: 1.64±0.02; 1.62–1.66). Promesonotum markedly convex, posterior region of mesonotum and propodeal dorsum approximately straight, with metanotal groove visible as suture; propodeal dorsum 3 × as long as declivity, their junction forming blunt angle. Anterior margin of petiolar node ca. 1/2 height of posterior margin and length of dorsal margin, separated from them by an angle and a convexity, respectively.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head present; near posterior margin of head with two elongate, erect hairs; antennal scape covered with erect hairs and lacking appressed hairs; junction of propodeal dorsum and declivity with two pairs of erect hairs.
Major worker. Unknown.
Camponotus aina is endemic to Madagascar, its distribution restricted to the dry forest and Tsingy of the RS Ankarana in the north of the island (Fig.
Camponotus aina A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217291 D distribution map.
Camponotus aina looks similar to C. joany in that it has a straight propodeal dorsum, but in the latter species the propodeal dorsum is 2 × as long as the declivity and the dorsal margin of the petiolar node is shorter than the posterior margin.
The species name aina is a non-Latin singular noun and is being used as a noun in apposition.
Madagascar: Province Mahajanga: PN de Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, dry forest, under stone, 08 Nov 2002 (Fisher, Griswold et al.) collection code: BLF06542, specimen code: CASENT0489917 (
2 minor workers and 1 major worker with same data as holotype but with specimen codes: CASENT0837586, CASENT0837587, CASENT0489918 (
Madagascar: Mahajanga: PN Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (
With head in full-face view, lateral margins of head anterior to eye level parallel, lacking erect hairs, lateral cephalic margin rounding to posterior margin, anteromedian clypeal margin continuously forming broad convexity; propodeal dorsum immediately posterior to metanotal groove convex, then concave medially and rounding to declivity surface.
Minor worker. In full-face view, head widest at midlength; lateral margins posterior to level of eye, rounding evenly to posterior margin; eye protruding and large (EL/CS: 0.25±0.01; 0.23–0.26), not breaking lateral cephalic margin; level of its posterior border located at ca. posterior 1/3 of head (PoOc/CL: 0.28±0.01; 0.27–0.30); frontal carinae posteriorly parallel (FR/CS: 0.25±0.01; 0.24–0.27); clypeus without anterolateral angle, its anteromedian margin broadly convex; mandible with six teeth, its two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.59±0.08; 1.46–1.71). Mesosoma long and low (MPH/ML: 0.28±0.01; 0.26–0.30), promesonotum weakly convex, mesonotum with posterior portion flat immediately anterior to a weakly visible metanotal groove; propodeal dorsum anteriorly convex, with feeble concavity medially, dorsal margin of propodeum rounding to declivity; propodeal dorsum 2 × as long as declivity. Petiolar node nodiform; with dorsal margin inclined posteriorly and forming a blunt angle to anterior face; anterior face of petiolar node 1/2 height of posterior face; femur of hind leg rounded axially and twisted basally.
First and second gastral tergites with a pair of white spots; erect hairs on lateral margin of head absent; two erect hairs present near posterior margin of head; antennal scape only covered with appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 3.10±0.18; 2.82–3.29; CWb/CL: 0.92±0.01; 0.90–0.94) with markedly concave posterior margin; apical 1/3 of antennal scape extending beyond posterior cephalic margin; robust mesosoma with propodeal dorsum slightly concave and its length ca. 2 × height of declivity; petiole tapering dorsally.
Geographically restricted to the PN Namoroka in the western dry forest of Madagascar (Fig.
Camponotus aro A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0489917 D distribution map.
Camponotus aro is morphologically similar to C. gouldi in that both species are characterized by a broadly convex anteromedian margin of clypeus, but the latter species is diagnosed as having a head that extends posteriorly into a broad, short neck and its propodeal dorsum is straight.
The definition for C. aro is supported by the congruence between the results of traditional qualitative morphology and the NC-clustering technique. The classification of the samples for the species is at 100% success.
The species name aro is a non-Latin singular noun used in apposition.
Madagascar: Province Toliara: PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest, ex rotten log, 05–09 Feb 2003 (Fisher, Griswold et al.) collection code: BLF07523, specimen code: CASENT0493252 (
3 workers, same data as holotype but with the following specimen codes: CASENT0493251, CASENT0837576, CASENT0837577 (
Madagascar: Toliara: 15 km E Sakaraha, -22.9, 44.68333, 760 m, tropical dry forest (P.S. Ward) (
With head in full-face view, lateral margins of head anterior to eye level, parallel, and lacking erect hairs; lateral cephalic margin rounding to posterior margin; anteromedian clypeal margin continuously forming broad convexity; propodeal dorsum more or less straight and separated from declivity surface by broad angle.
Minor worker. In full-face view, head sides anterior to level of eye parallel, converging progressively to posterior margin behind eye level; eyes protruding and large (EL/CL: 0.26±0.01; 0.23–0.28), breaking lateral cephalic margin, level of its posterior border approximately located at posterior 1/4 of head (PoOc/CS: 0.26±0.01; 0.24–0.28); frontal carinae posteriorly parallel (FR/CS: 0.26±0.01; 0.23–0.27); clypeus without well-defined anterolateral angle, its anteromedian margin broadly convex; mandible with six teeth, the two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.57±0.08; 1.38–1.68). Promesonotum weakly convex, mesopropodeum almost flat; mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum approximately straight, rounding progressively towards declivity; propodeal declivity ca. 1/3 length of dorsum. Petiolar node flattened anteroposteriorly or short and high, tapering dorsally; femur of hind leg flattened laterally and twisted near base.
First and second gastral tergites with a pair of white spots; erect hairs lacking on lateral margin of head; posterior margin of head with a pair of erect hairs; antennal scape only covered with appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum without erect hairs.
Major worker. With characteristics of minor worker, except for the following characters: enlarged head (CS: 3.22±0.15; 3.03–3.39; CWb/CL: 0.93±0.01; 0.92–0.93) with noticeable medial excision on posterior margin; straight anterior clypeal margin; apical 1/4 of antennal scape surpassing posterior cephalic margin; propodeum dorsum and declivity the same length.
Camponotus asara is endemic to Madagascar and geographically restricted to the dry forest of the PN Zombitse in the southern high plateau of the island and the relict montane rainforest of the PN Ambohijanahary (Fig.
Camponotus asara A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0493252 D distribution map.
Camponotus asara is morphologically similar to C. bozaka and C. becki in that its petiolar node is more or less compressed anteroposteriorly in lateral view and its body color is black to dark brown, or the posterior portion of the mesosoma is pale brown to yellow. However, C. bozaka has a scalelike petiolar node, short and high mesosoma, and propodeal dorsum that joins the declivity in a blunt angle with a length < 2 × height of the declivity, the junction being angulate. Regarding C. becki, its anteromedian clypeal margin is truncate and a few erect hairs are present on the lateral margin of the head behind the level of the posterior ocular margin.
The cluster of the samples for C. asara in the NC-clustering dendrogram is supported by LDA with a classification success of 100%.
The species name asara is a non-Latin singular noun used in apposition.
Madagascar: Province Toliara: 3.5 km 236° SW Marovato, -25.55389, 45.25583, 230 m, spiny forest/thicket, ex rotten log, 14 Feb 2002 (Fisher, Griswold, and Arthropod Team) collection code: BLF05595, specimen code: CASENT0454042 (
1 major worker same data as holotype but specimen coded as: CASENT0454043 (major) (
Madagascar: Antsiranana: Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest, (J.-J. Rafanomezantsoa et al.) (
With head in full-face view, lateral margins of head anterior to eye level parallel, lacking erect hairs; in oblique profile, four or more pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin; clypeus with distinct anterolateral corner; in profile, junction of propodeal dorsum to declivity rounded; petiole nodelike and not anteroposteriorly compressed.
Minor worker. With head in full-face view, lateral margins anterior to eye level parallel, converging abruptly towards posterior margin behind eye level; eye large and convex (EL/CS: 0.28±0.01; 0.26–0.30), interrupting lateral cephalic border, level of its posterior margin situated approximately at posterior 1/3 of head (PoOc/CL; 0.27±0.01; 0.25–0.29); frontal carinae close to each other, their distance equal to or smaller than their smallest distance to eye (FR/CS: 0.23±0.01; 0.22–0.25); clypeus with anterolateral angle and broadly convex anteromedian margin; mandible with two apical teeth distant from each other; antennal scape relatively long (SL/CS: 1.66±0.09; 1.48–1.84). Promesonotum weakly convex and mesopropodeum feebly convex (MPH/ML: 0.33±0.02; 0.29–0.36), mesonotum flat near weakly visible metanotal groove; propodeal dorsum anteriorly convex, posteriorly flat, rounding to declivity; propodeal dorsum 2 × as long as declivity. Petiole nodiform, its dorsal margin inclined posteriorly, rounding to anterior margin; anterior face 1/2 height of posterior face; femur of hind leg rounded axially, not twisted near base.
First and second gastral tergites without a pair of white spots; lateral margin of head without erect hairs; posterior cephalic margin with a pair of erect hairs; in profile, four pairs of erect hairs arranged from level of anterior margin of eye to posterior cephalic margin; erect hairs lacking from antennal scape, pubescence present; pronotum with a pair of erect hairs; posterodorsal angle of propodeum without erect hairs.
Major worker. With characteristics of minor worker, except for the typically broader head (CS: 3.23±0.34; 2.80–3.77; CWb/CL: 0.97±0.04; 0.89–1.03), with broadly concave posterior margin; apical 1/3 of antennal scape extending beyond posterior cephalic margin; robust mesosoma, with propodeal dorsum convex immediately posterior to metanotal groove and < 2 × as long as declivity; petiolar node compressed anteroposteriorly.
Camponotus atimo, endemic to Madagascar, generally occurs in the southern part of the island (Fig.
Camponotus atimo A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0454042 D distribution map.
Camponotus atimo may be difficult to distinguish from C. tapia in that both species have the same body shape and the dorsum of the head is fringed with four or more pairs of erect hairs arranged successively from the level of the anterior ocular margin towards the posterior cephalic margin. However, in C. tapia, the junction of the propodeal dorsum to declivity surface is angulate.
Camponotus atimo also may be confounded with C. strangulatus, but the latter is characterized by the head having only three pairs of erect hairs arranged successively from the level of anterior ocular margin towards the posterior cephalic margin.
The identity of C. atimo, based on traditional qualitative taxonomy, has been confirmed by multivariate morphometrics. The grouping of the samples of C. atimo generated by the NC-clustering method is corroborated by cumulative LDA with a classification success of 100%.
This new name atimo is a singular non-Latin noun used in apposition, derived from the Malagasy word for “south” in reference to the restricted distribution of the species to the southern half of Madagascar.
Camponotus aurosus
Roger, 1863: 134. Syntype workers, Mauritius (Roger) (ZMHB); one syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0104620 (ZMHB) [examined]. [Combination in Camponotus (Myrmepomis): Emery, 1920a: 258; in Camponotus (Myrmosericus): Forel, 1914: 268;
Mauritius: [Mauritius, 19161], -20.3, 57.583333, (Roger) (
In full-face view, lateral margin of head anterior to eye level diverging posteriorly and covered with erect hairs; anterior clypeal margin broadly triangular or convex; mesosoma in profile short and high; gastral tergites covered with abundant pubescence; head and gaster black, mesosoma reddish orange to brown.
Minor worker. In full-face view, lateral cephalic margins diverging posteriorly, widest at eye level, rounding evenly to posterior margin; eye protruding and large (EL/CS: 0.27±0.01; 0.25–0.29), breaking lateral margin of head, its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.24±0.01; 0.22–0.26); frontal carinae widely opened posteriorly (FR/CS: 0.30±0.01; 0.29–0.32), posteriorly diverging; clypeus without well-defined anterolateral angle, its anteromedian margin broadly convex or triangular; mandible with six teeth, the two apical teeth distant from each other; antennal scape relatively long (SL/CS: 1.33±0.04; 1.27–1.40). Dorsal outline of mesosoma strongly arched, metanotal groove weakly visible; propodeal dorsum almost straight, joining the declivity at a blunt angle; propodeal declivity 3/4 length of the dorsum; petiolar node short, high, and tapering dorsally, its dorsal margin inclined posteriorly and forming a blunt angle with anterior face; anterior face of petiolar node 1/2 height of posterior face; tibia of hind leg rounded axially, without basal twist.
First and second gastral tergites without a pair of white spots; lateral margin of head anterior to level of eyes with a few erect hairs, which are absent behind level of eyes; antennal scape without erect hairs but covered with appressed hairs; pronotum with a pair of erect hairs; posterior cephalic margin with three or more pairs of erect hairs; posterodorsal angle of propodeum with two pairs of erect hairs.
Major worker. Differing from minor worker in the following characters: larger head (CS: 2.24±0.11; 2.15–2.37; CWb/CL: 0.92±0.03; 0.87–0.95), with broadly slight medial concavity on posterior margin; apical 1/3 of antennal scape surpassing posterior cephalic margin; with mesosoma in lateral view, length of propodeal dorsum the same as height of propodeal declivity; petiolar node more compressed anteroposteriorly.
The distribution of C. aurosus is restricted to Mauritius and Reunion islands (Fig.
Camponotus aurosus A lateral view B head in full-face view C dorsal view of minor worker CASENT0064815 D distribution map.
Camponotus aurosus is similar to C. hagensii with respect to the shape of the head and the presence of erect hairs on the lateral cephalic margin, but the latter is characterized by the straight anteromedian margin of its clypeus, low and long mesosoma, and the presence of some pubescence on the gastral tergites.
The grouping of the C. aurosus samples in the same cluster shown by the dendrogram of multivariate morphometric analysis is corroborated by the cumulative LDA at 100% identification success. This result is congruent with that of the qualitative morphology-based study.
Camponotus radamae st. becki
Santschi, 1923: 292. Syntype workers, Madagascar (R. Beck) (
Madagascar: Fianarantsoa: PN Andringitra, Plateau d’Andohariana, 35.9 km 205° Ambalavao, -22.15233, 46.89917, 2000 m, ericoid thicket (B.L. Fisher et al.) (
With head in full-face view, lateral margins of head anterior to eye level parallel and lacking erect hairs, lateral cephalic margin rounding to posterior margin, anteromedian clypeal margin truncate; propodeal dorsum more or less straight and joining declivity surface at a broad angle; body color black to dark brown.
Minor worker. With head in full-face view, lateral margins anterior to level of eye parallel, rounding evenly to a straight posterior margin; eye more or less convex (EL/CS: 0.27±0.02; 0.25–0.30), not breaking lateral cephalic margin, level of its posterior margin located generally at posterior 1/4 of head (PoOc/CL: 0.26±0.01; 0.24–0.27); frontal carinae posteriorly parallel (FR/CS: 0.27±0.01; 0.26–0.29); clypeus without well-defined anterolateral angle and with anteromedian margin broadly convex; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 0.27±0.01; 0.26–0.29). Promesonotum weakly convex, mesopropodeum generally flat; mesonotum with posterior portion flat immediately anterior to a weakly visible metanotal groove; propodeal dorsum anteriorly convex and posteriorly flat, joining declivity blunt angle; propodeal dorsum 2 × as long as declivity. Petiolar node short and high, its dorsal margin rounding to anterior face, height of its anterior face 2/3 that of posterior face; femur of hind leg flattened laterally, twisted near base.
First and second gastral tergites usually without white spots; lateral margin of head without erect hairs; two erect hairs near posterior margin of head; antennal scape only covered with appressed hairs. Pronotum with a pair of erect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. With characteristics of minor worker except: enlarged head (CS: 2.80±0.23; 2.54–2.96; CWb/CL: 0.97±0.03; 0.94–1.00) with slight, medially concave posterior margin; apical 1/3 of antennal scape surpassing posterior cephalic margin; robust mesosoma, metanotum distinctly visible; propodeal dorsum same length as declivity; petiolar node flattened anteroposteriorly.
Based on recent collection data, Camponotus becki is restricted to the ericoid thickets and montane rainforests of the PN Andringitra (Fig.
Camponotus becki A lateral view B head in full-face view C dorsal view of minor worker CASENT0071380 D distribution map.
See discussion under C. asara. The taxonomic categorization of C. becki based on the study of qualitative morphology is supported by the NC-clustering method. This species is identified by LDA at 100% success.
Madagascar: Province Toliara: PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest, pitfall trap (Fisher, Griswold et al.) 05–09 Feb 2003, collection code: BLF07507, specimen code: CASENT0050097 (
Madagascar: Antsiranana: RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (
In full-face view, lateral cephalic margins converge posteriorly towards eye level and without erect hairs posterior to eye level; two apical teeth of mandible positioned close to each other.
Minor worker. In full-face view, head sides converging progressively towards posterior margin; eye convex and large (EL/CL: 0.30±0.01; 0.29–0.34), breaking lateral cephalic borders, level of its posterior margin generally situated at posterior 1/3 to 1/4 of head (PoOc/CL: 0.26±0.01; 0.24–0.28); frontal carinae posteriorly parallel to each other (FR/CS: 0.26±0.01 0.25–0.29); clypeus without well-defined anterolateral angle and with broadly convex anteromedian margin; mandible with six teeth, of which the two apical are close together; antennal scape relatively long (SL/CS: 1.63±0.10; 1.47–1.87) with suberect hairs inclined 30° and pubescence. Promesonotum weakly convex, mesopropodeum almost flat, mesonotum flat immediately anterior to metanotal groove; metanotal groove weakly visible; propodeal dorsum anteriorly convex and posteriorly flat; junction of dorsal margin to declivity rounded; propodeal declivity height 1/2 length of propodeal dorsum; petiole nodiform, its dorsal margin rounding to anterior margin; anterior face 1/2 height of posterior face; tibia of hind leg rounded, not twisted basally.
First and second gastral tergites without a pair of white spots; erect hairs covering lateral margin of head anterior to eye level but absent behind eyes; posterior margin of head with more than six erect hairs; antennal scape with suberect hairs inclined at ca. 30° and pubescence; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Unknown
Camponotus bemaheva is restricted to the spiny bush and thicket of the southern region and the dry forest habitats of western Madagascar (Fig.
Camponotus bemaheva A lateral view B head in full-face view C dorsal view of minor worker CASENT0154158 D distribution map.
Workers of C. bemaheva look very similar to those of C. daraina in that they have closely spaced apical teeth on the mandibles and the lateral margins of their head converge posteriorly towards eye level, but the latter is characterized by the presence of erect hairs on the lateral cephalic margin posterior to eye level.
The alignment of C. bemaheva in the same cluster displayed by the dendrogram of multivariate morphometric method is confirmed by the cumulative LDA at 100% identification success, validating the taxonomy hypothesized by qualitative morphology.
The new species bemaheva is a non-Latin proper noun used in apposition and is named in honor of Fidelis Bemaheva for his help collecting ants in the Malagasy region.
Camponotus maculatus r. boivini
Forel, 1891: 34. Syntype workers, queen and male, Madagascar (Sikora) (
Camponotus maculatus st. fairmairei
Santschi, 1911a: 130. Syntype workers Madagascar, colony living with larvae of Fulgorids (Fairmaire, 1900) (
One major worker and one male specimen that are labeled respectively with CASENT0104638 and CASENT0104637 (ZMHB), were collected from Mahajanga, and are labeled with an unpublished name “Camponotus maculatus subsp. hova var. laticollus” by Forel. In the present study, the two specimens are identified as Camponotus boivini.
Madagascar: Antananarivo: Alasora, -18.96245, 47.58925, 1434 m, eucalyptus plantation (B.L. Fisher et al.) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible normally spaced; antennal scape covered with suberect hairs inclined at ca. 45°; in lateral view, posterior 1/2 of mesonotum to posterodorsal corner of propodeum somewhat convex, propodeal dorsum ca. 2 × as long as the height of declivity surface.
Minor worker. In full-face view, lateral cephalic borders anterior to level of eye parallel to each other, converging progressively towards posterior margin behind eye level; eye protruding and large (EL/CS: 0.32±0.02; 0.29–0.36), breaking lateral cephalic margins; head sides behind eye level 1/4 length of head (PoOc/CL: 0.27±0.01; 0.24–0.30); frontal carinae widely diverging posteriorly (FR/CS: 0.32±0.01; 0.30–0.35); clypeus with anterolateral angle, its anteromedian margin with blunt angle or convex; mandible with two apical teeth distant from each other; antennal scape relatively long (SL/CS: 1.37±0.10; 1.22–1.56). Promesonotum weakly convex and mesopropodeum feebly convex, mesonotum with posterior portion flat immediately anterior to metanotal groove, metanotum weakly visible, propodeal dorsum anteriorly convex and posteriorly flat, dorsal margin of propodeum and declivity joining at a blunt angle, height of propodeal declivity 1/2 length of propodeal dorsum. Petiolar node flattened, short, and high, without noticeable dorsal margin; tibia of hind leg rounded axially, without twist in the basal portion.
First and second gastral tergites without a pair of white spots; lateral margin of head anterior to eyes level with erect hairs, which are absent behind eyes; posterior margin of head with a pair of erect hairs; antennal scape covered with erect hairs inclined at 45°; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in having larger head (CS: 2.29±0.28; 1.96–3.12; CWb/CL: 0.91±0.04; 0.86–0.99), short antennal scape barely surpassing posterior cephalic margin; robust mesosoma with distinct metanotum, and petiolar node as high as long. Other characters as in minor worker.
The distribution of C. boivini is generally limited to western and the high plateau of Madagascar (Fig.
Camponotus boivini A lateral view B head in full-face view C dorsal view of minor worker CASENT0486575 D distribution map.
Camponotus boivini may be confused with C. cemeryi but the latter is characterized by a short and high mesosoma which shows a strongly convex dorsal outline. Camponotus boivini can be confounded with C. mixtellus but workers of the latter species have an antennal scape covered with suberect hairs inclined at ca. 30°, are generally larger (CS: 1.56±0.13; 1.16–1.83; ML: 2.76±0.16; 3.10–3.45), and are mostly found in the rainforest of the region.
The definition of C. boivinii based on traditional qualitative morphology is congruent with the results achieved by the exploratory analysis and the cumulative LDA, which has an identification success of 100%.
Madagascar: Province Fianarantsoa: Parc National d’Andringitra, Plateau d’Andohariana, 39.8 km 204° Ambalavao, -22.18767, 46.90083, 2150 m, rubicole thicket at base of cliff, under stone, 16 Apr 2006 (B.L. Fisher et al.) collection code: BLF13773, specimen code: CASENT0217309 (
1 major worker with same data as holotype but with specimen code CASENT0071370 (
Madagascar: Fianarantsoa: Manandriana I Non Protected Area, 27.11 km SW Ambositra, -20.73194, 47.09413, 1590 m, Savannah grassland (A. Ravelomanana) (
With head in full-face view, lateral margin of head anterior to eye level parallel, lacking erect hairs; clypeus with noticeable anterolateral corner; in profile, petiolar node anteroposteriorly compressed.
Minor worker. In full-face view, lateral cephalic margins anterior to level of eye approximately parallel, evenly rounding to posterior margin; eye protruding and large (EL/CS: 0.28±0.01; 0.26–0.30), breaking lateral cephalic margin, level of its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.25±0.01; 0.23–0.26); frontal carinae close to each other, their distance equal to or smaller than smallest distance of one to the eye (FR/CS: 0.28±0.01; 0.28–0.29); clypeus with anterolateral corners, its anteromedian margin broadly triangular or convex; mandible with two apical teeth normally spaced; antennal scape relatively short (SL/CS: 1.19±0.05; 1.08–1.25). Promesonotum weakly convex, mesopropodeum almost flat; mesonotum flat immediately anterior to metanotal groove; metanotal groove weakly visible; dorsal margin of propodeum straight and joining declivity at a noticeable angle, propodeal dorsum < 2 × as long as declivity. Petiolar node scalelike with sharp dorsal margin; femur of hind leg rounded axially and not twisted basally.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head not present; posterior margin of head with two erect hairs; with head in profile, four pairs of erect hairs arranged from level of anterior margin of eye to posterior cephalic margin; antennal scape covered only with appressed hairs; pronotum, mesonotum, and posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 2.07±0.18; 1.91–2.40; CWb/CL: 0.95±0.03; 0.90–0.98) with slightly concave to almost straight posterior margin; apical 1/3 of antennal scape extending beyond posterior cephalic margin; robust mesosoma with length of propodeal dorsum approximately the same as height of declivity.
Known only from the south-central high plateau of Madagascar, C. bozaka occupies rubicole thicket, Uapaca woodland, savannah grassland, and shrubland habitats (Fig.
Camponotus bozaka A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217309 D distribution map.
See discussion under C. asara. The taxonomic identity of C. bozaka based on conventional qualitative morphology is supported by multivariate morphometric analysis. The grouping shown by the morphometric dendrogram and confirmed by cumulative LDA at 100% success supports the existence of the species.
This new name bozaka is a singular non-Latin noun used in apposition and is derived from the Malagasy word for “dried grass” in reference to the abundant grassland where this species occurs.
Camponotus hova cemeryi
Özdikmen, 2010: 34. Syntype major workers, queen and male, Madagascar, Majunga (Voeltzkow); 1 syntype major worker designated as lectotype, by present designation, Madagascar, Majunga (Voeltzkow) AntWeb CASENT0101103 (
Camponotus hova var. luteolus Emery, is a junior secondary homonym of Camponotus maculatus luteolus Emery, 1906:188. As a subspecies of Camponotus bonariensis luteolus Emery, 1920a: 233; [First available use of Camponotus maculatus r. hova var. luteolus Forel, 1897: 187; unavailable name].
Madagascar: Antananarivo: Navoatra VI Non Protected Area, 7.3 km NW Arivonimamo, -18.97889, 47.12253, 1276 m, Uapaca woodland (A. Ravelomanana) (
With head in full-face view, lateral cephalic margins anterior to eye level parallel and lacking erect hairs, lateral and anteromedian clypeal margin continuously forming broad convexity; scape covered with erect hairs; mesosoma short and high (MPH/ML: 0.39, 0.46); petiolar node flattened anteroposteriorly; body color yellowish to brown.
Minor worker. In full-face view, head sides anterior to level of eye parallel, converging progressively to posterior margin; eye large and convex (EL/CS: 0.33±0.01; 0.31–0.35), breaking lateral cephalic margin, level of its posterior border generally located from posterior 1/3 to 1/4 of head (PoOc/CL: 0.25±0.02; 0.22–0.29); frontal carinae not strongly diverging posteriorly (FR/CS: 0.28±0.01; 0.26–0.30), posteriorly diverging; clypeus with anterolateral angle, its anteromedian margin with blunt or convex angle, two apical teeth of mandible distantly spaced; antennal scape long (SL/CS: 1.49±0.05; 1.41–1.60). Dorsum of mesosoma strongly arched (MPH/ML: 0.42±0.02; 0.39–0.46), outline of promesonotum and dorsum of propodeum evenly convex; metanotal groove weakly visible, dorsal margin and declivity of propodeum connect at blunt angle; propodeal declivity height > 3/4 of dorsum length; petiolar node flattened anteroposteriorly, without noticeable dorsal margin; tibia of hind leg rounded axially and not twisted basally.
First and second gastral tergites without a pair of white spots; whole lateral margin of head covered with erect hairs; posterior margin of head with two pairs of erect hairs; antennal scape covered only with appressed hairs; posterodorsal angle of propodeum with more than four erect hairs.
Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 2.41±0.22; 2.19–2.72; CWb/CL: 0.87±0.05; 0.83–0.94); stronger mandible; apical 1/5 of antennal scape surpassing posterior cephalic margin; metanotum visible; propodeal dorsum slightly convex joining a vertically built declivity at rounded angle; petiolar node much higher than long.
Camponotus cemeryi occurs in dry forest and spiny forest of west Madagascar and in the savannah shrubland and woodland of the central plateau (Fig.
Camponotus cemeryi A lateral view B head in full-face view C dorsal view of minor worker CASENT0217306 D distribution map.
Members of C. cemeryi can be separated from similar species like C. boivini by the strongly convex dorsal outline of its short and high mesosoma. It can be differentiated from C. mahafaly by the presence of erect to suberect hairs on the antennal scape.
Worker specimens that show the typical form of C. cemeryi are generally found across the western dry forest of Madagascar, but specimens have been collected from the mountaintops on the high plateau of the island that present morphological variation in which the lateral margin of head posterior to eye level is covered with erect hairs, three pairs of erect hairs are present on posterior cephalic margin, and body color is generally depigmented yellow or brown with depigmented yellow in some parts of the body. Based on these characters, this variant may constitute a separate species; however, the dendrogram based on quantitative morphological analysis did not support this hypothesis. The variant is nested within the cluster of the typical form of the species according to the NC-clustering method, and all of these samples were correctly classified by LDA at 100% success.
Camponotus cervicalis
Roger, 1863: 134. Syntype workers, Madagascar (Humboldt) (
Camponotus gaullei
Santschi, 1911a: 128. Syntype workers, Madagascar, S. de la Baie d’Antongil, (
Camponotus perroti
Forel, 1897: 202. Syntype workers, Madagascar, Sainte Marie (Perrot) (
Camponotus perroti aeschylus
Forel, 1913: 224. Holotype (by monotypy) alate queen, Madagascar (C. Keller) AntWeb CASENT0101561 (
Camponotus gerberti
Donisthorpe, 1949: 271. Syntype workers, Madagascar, Sainte Marie (R. Géberti 1917) (
Madagascar: Antananarivo: [Madagascar]; Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (Humblot) (
With head in full-face view, lateral cephalic margin converging posteriorly towards eye level, covered with erect hairs from anterior to posterior of eye level; anteromedian margin of clypeus broadly convex; two apical teeth of mandible normally spaced.
Minor worker. In full-face view, head sides converging progressively towards short posterior margin; eye protruding and large (EL/CS: 0.28±0.01; 0.26–0.30), breaking lateral cephalic margin, level of posterior margin located at posterior 1/3 of head (PoOc/CL: 0.30±0.01; 0.29–0.31); frontal carinae wide (FR/CS: 0.27±0.01; 0.26–0.28), distance between them larger than smallest distance to eye; clypeus with anterolateral angle and anteromedian margin with blunt angle or convex; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.81±0.12; 1.57–1.94). Promesonotum weakly convex, mesopropodeum almost flat, joining declivity with rounded angle; metanotal groove weakly visible; propodeal declivity 1/2 length of the dorsum. Petiolar node as long as high, with dorsal margin inclined posteriorly and rounding to anterior margin; anterior face 1/2 height of the posterior; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; lateral margin of head with erect hairs; more than six erect hairs present near posterior margin of head; antennal scape covered with erect to suberect hairs inclined at ca. 45° as well as appressed hairs; pronotum covered with few erect hairs, mesonotum with a pair of hairs; posterodorsal angle of propodeum with two pairs of erect hairs.
Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 4.17±0.35; 3.83–4.63; CWb/CL: 0.91±0.03; 0.85–0.93); the more strongly built mandible; apical 1/4 of antennal scape surpassing posterior cephalic margin; pronotum convex, mesonotum sloping towards metanotum, propodeal dorsum feebly convex and its junction to declivity broadly angulate; petiolar node much higher than long.
Camponotus cervicalis is only known to occur in littoral forests and lower elevations of the rainforests in the east of Madagascar between the Ambanitaza forest in the north and the RNI Betampona in the south (Fig.
Camponotus cervicalis A lateral view B head in full-face view C dorsal view of minor worker CASENT0217303 D distribution map.
Camponotus cervicalis and C. niavo look similar, but in the latter, the level of the posterior ocular margin is located at posterior 1/4 of length of head and its antennal scape lacks appressed hairs.
In his original description of Camponotus gaullei,
The description of C. perroti by
Forel’s description of C. perroti aeschylus was based only on a single alate queen, and the comparison of this specimen with the queen specimens of C. cervicalis collected from its geographical range across Madagascar revealed that this type specimen was a queen of C. cervicalis.
Donisthorpe’s description of Camponotus gerberti did not compare the characters of the taxa to those of other species, although this description morphologically matches the type specimens of C. cervicalis. The data obtained from the recent ant survey in Madagascar and this information are sufficient to reasonably synonymize C. gerberti under C. cervicalis.
The identity of C. cervicalis based on conventional qualitative taxonomy is recognized by multivariate morphometric analysis. The cluster of the samples of the species created by NC-clustering method is confirmed by cumulative LDA with an identification success of 100%.
Madagascar: Province Antsiranana: Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 725 m, rainforest, ex rotten log, 03 Dec 2003 (B.L. Fisher et al.) collection code: BLF09678, specimen code: CASENT0077433 (
1 major worker of same data as holotype but with specimen code: CASENT0809930 and 5 minor workers of collection code BLF09664 and the following specimen codes: CASENT0077671, CASENT0835653, CASENT0077673, CASENT0077670, CASENT0077673 (
Madagascar: Antsiranana: Forêt Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (
In full-face view, lateral cephalic margins converge posteriorly towards eye level, area posterior to eye level covered with erect hairs; two apical teeth of mandible closely spaced; body color reddish orange.
Minor worker. With head in full-face view, lateral borders gradually converge to anterior eye level, strongly converging behind eye level; eyes protruding and large (EL/CS: 0.28±0.01; 0.26–0.30), interrupting lateral cephalic border, level of its anterior margin located at posterior 1/3 of head (PoOc/CL: 0.29±0.01; 0.28–0.30); frontal carinae not widely opened posteriorly (FR/CS: 0.25±0.001; 0.24, 0.26), distance between them larger than smallest distance to eye; clypeus without well-defined anterolateral angle and with anteromedian margin broadly convex; mandible with two apical teeth closely placed; antennal scape relatively long (SL/CS: 1.72±0.06; 1.56–1.82). Promesonotum weakly convex, mesonotum posterior portion flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight, rounding to short declivity 1/2 length of propodeal dorsum. Petiolar node is short and high, with dorsal margin inclining posteriorly and forming a blunt angle with anterior face that has a height ca. 1/2 that of posterior face; femur of hind leg axially rounded, not twisted near base.
First and second gastral tergites without a pair of white spots; head covered with erect hairs on lateral margin, six erect hairs near its posterior margin; antennal scape covered with suberect hairs inclined ca. 30°; pronotum and anterior part of mesonotum covered with erect hairs; posterodorsal angle of propodeum with two pairs of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 13.54±0.25; 3.12–3.76; CWb/CL: 0.90±0.04; 0.85–0.94) with markedly concave posterior margin; apical 1/4 of antennal scape surpassing posterior margin of head; two apical teeth of mandible distantly spaced; robust mesosoma, with promesonotum forming even convexity, metanotum distinctly visible, propodeal dorsum slightly convex immediately behind metanotum, joining declivity surface at a blunt angle, < 2 × as long as declivity; petiolar node much higher and more compressed anteroposteriorly.
The distribution of C. daraina is generally limited to the northern and western slopes of Madagascar. The species occurs in dry forest habitats, transitional forests, and rainforests of the Daraina region, transitional forests of Montagne d’Ambre and the Ampasindava Peninsula, and rainforests of RS Manongarivo (Fig.
Camponotus daraina A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0077433 D distribution map.
Camponotus daraina looks similar to C. bemaheva and C. roeseli but C. bemaheva lacks erect hairs from the lateral margin of the head posterior to the eye level and in C. roeseli the integument is either entirely reddish black or the head and the gaster are reddish black and the mesosoma is yellowish orange to reddish orange.
The identification of C. daraina based on conventional morphology-based taxonomy has been confirmed by multivariate morphometrics. The grouping of the samples of C. daraina produced by the NC-clustering method is corroborated by LDA with a classification success of 100%.
The species name daraina is a singular non-Latin noun used in apposition and refers to the type locality.
Camponotus dufouri
Forel, 1891: 16, 74 (Key). Syntype workers, Madagascar, 30 miles NW Tamatave (O’Swald) (
Camponotus dufouri imerinensis
Forel, 1891: 18. Syntype workers and queen, Madagascar, Environ d’Antananarivo, Imerina [labeled as “Andrangoloaka”] (Sikora) (
Madagascar: Antananarivo: [Madagascar], Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (Sicora) (
With head in full-face view, lateral cephalic margins converge posteriorly towards eye level, strongly converging from posterior ocular margin to occipital corner of head; anteromedian margin of clypeus straight; two apical teeth of mandible normally spaced; lateral cephalic margin anterior to eye level covered with erect hairs.
Minor worker. In full-face view, lateral cephalic margins converging progressively towards posterior margin or slightly converging to level of anterior ocular margin and strongly converging posteriorly; eye large and convex (EL/CS: 0.28±0.02; 0.25–0.31), breaking lateral cephalic border, located at midlength of head, level of its posterior margin within the posterior 1/3 of head (PoOc/CL; 0.33±0.01; 0.31–0.36); frontal carinae more or less wide, posteriorly parallel (FR/CS: 0.25±0.01; 0.24–0.27), their distance larger than smallest distance to eye; clypeus with anterolateral angle and medially slightly concave anteromedian margin; two apical teeth of mandible distantly spaced; antennal scape relatively long (SL/CS: 1.93±0.08; 1.76–2.12). Mesosoma long and low (MPH/ML: 0.29±0.01; 0.27–0.32), with slightly convex promesonotum, nearly flat mesopropodeum, and feebly visible metanotal groove; junction of propodeal dorsum to declivity bluntly angulate; propodeal declivity 1/3 length of dorsum. Petiole nodiform, tapering dorsally; its dorsal margin inclined posteriorly and forming a blunt angle to anterior face; posterior face 3 × as high as the anterior; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; lateral margin of head covered with erect hairs; near posterior margin of head with more than six erect hairs; antennal scape covered with erect hairs inclined at ca. 45°; pronotum with numerous erect to suberect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. With characteristics of minor worker except for the following divergent characters: larger head (CS: 3.93±0.51; 3.46–4.54; CWb/CL: 0.88±0.08; 0.80–0.97), with concavity on posterior margin; anteromedian margin of clypeus slightly concave; apical 1/3 of antennal scape surpassing posterior cephalic margin; promesonotum and propodeum in separate convexity; mesonotum and propodeal dorsum 2 × as long as declivity, with blunt angle junction.
Camponotus dufouri is only known from Madagascar. Its distribution follows that of eastern humid forests ranging from the littoral to the montane rainforest of the island (Fig.
Camponotus dufouri A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT0060837 and CASENT0274127 G distribution map.
A clypeus with a straight anteromedian margin makes C. dufouri easy to separate from C. lokobe. Also, in C. tendryi, when the head is set in full-face view, the lateral cephalic margin posterior to the eye level is without erect hairs and its body size is smaller (CS: 1.57±0.01; 1.56–1.58), while in C. dufouri, erect hairs are present on the lateral cephalic margin posterior to the eye level and its body size is larger (CS: 1.97±0.19; 1.66–2.26).
A verification of the syntype workers of C. dufouri imerinensis demonstrates that they were indistinguishable from the syntypes of C. dufouri. Moreover, since the geographic distribution of C. dufouri is ranging from the littoral to the montane rainforests of Madagascar where the type specimens of the subspecies name were collected, it is consequently correct to synonymize it under C. dufouri.
The qualitative morphology-based study of C. dufouri agrees with the multivariate statistical analysis and both methods support the taxonomic delimitation for this species. The validity of the species, supported by the grouping obtained from NC-clustering, is confirmed by LDA with an identification success of 100%.
Camponotus quadrimaculatus var. gibber
Forel, 1891: 59. Lectotype minor worker, by present designation, Madagascar, Andrangoloaka (Sikora) AntWeb CASENT0101513 (
Madagascar: Antananarivo: [(de Diversa); Museum Paris, Grandidier 1899]; Mantasoa; Manjakandriana, -19.033333, 47.9166666, 1409 m (
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; two pairs of white spots present on second and third abdominal tergites; pronotum, mesonotum, and propodeum forming separate convexities, metanotal groove depressed; level of propodeum lower than that of promesonotum.
Minor worker. In full-face view, head sides diverging towards broadly convex posterior margin; eye slightly protruding and small (EL/CS: 0.27±0.01; 0.24–0.30), not breaking lateral cephalic margin, level of its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.25±0.01; 0.23–0.28); frontal carinae wide and diverging posteriorly (FR/CS: 0.35±0.01; 0.33–0.36), distance between them larger than their smallest distance to eye; clypeus with anterolateral angle and straight anteromedian margin; mandible with two apical teeth distant from each other; antennal scape relatively long (SL/CS: 1.17±0.07; 0.93–1.27). Pronotum and mesonotum forming a separate convexity; propodeal dorsum convex anteriorly, concave medially, then flat posteriorly, joining declivity in noticeable angle; metanotal groove weakly visible; propodeal declivity 3/4 length of dorsum. Petiolar node short and high, with dorsal margin straight then rounding to both anterior and posterior faces; anterior face almost 2/3 height of posterior face; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites with a pair of white spots; lateral margin of head without erect hairs; three pairs of erect hairs present near posterior margin of head; antennal scape only covered with appressed hairs; pronotum with few erect hairs; mesonotum with a pair of erect hairs; posterodorsal corner of propodeum with two pairs of erect hairs. Body color shining brown to dark brown; apical section of appendages lighter in color.
Major worker. With characteristics of minor worker except: enlarged head (CS: 2.45±0.21; 2.21–2.82; CWb/CL: 1.05±0.04; 1.01–1.10) with broadly concave posterior margin; anteromedian clypeal margin noticeably excised medially; antennal scape hardly extending beyond posterior cephalic margin; robust mesosoma, pronotum, and mesonotum an even convexity, metanotum distinct, propodeal dorsum sloping straight to declivity, approximately the same length as declivity; petiolar node more flattened anteroposteriorly.
Camponotus gibber occurs in mid-altitude rainforest, montane rainforest, open areas on the forest edge, and the savannah grassland of the high plateau of Madagascar. Its distribution ranges from the RS Manongarivo and PN Marojejy in the north through the Corridor Forestier Analamay-Mantadia and PN Andringitra in the south-central region to the PN Andohahela and Anosyenne Mountains in the south. It has also colonized the littoral rainforest of Ambondrobe Vohemar, the dry forest of the PN Isalo, and cultivated land in the Andrambovato Forest (Fig.
Camponotus gibber A lateral view B head in full-face view C dorsal view of minor worker CASENT0188619 D distribution map.
Camponotus gibber may be difficult to differentiate from C. rotrae and C. quadrimaculatus in that they have two pairs of white spots on the second and third abdominal tergites. In both latter species, however, the pronotum, mesonotum, and propodeum do not form separate convexities, the metanotal groove is not depressed, and the dorsal face of its petiolar node joins the posterior face at an angle.
There are apparently three forms within C. gibber. These are geographically isolated across their distribution along the eastern rainforest of Madagascar due to the presence of high mountain chains in northwestern Madagascar. In the first form, the mesosoma strongly forms separate convexities and the propodeal dorsum is more or less straight. The second form is characterized by a more or less continuous dorsal outline of the mesosoma and a broadly concave propodeal dorsum. The third form constitutes intermediate degrees of these phenotypic variations because workers present separate convexities of mesosoma and a slightly concave propodeal dorsum. The members of the first two forms show morphological variabilities that gradually merge in the third form.
The relatively low 91.89% classification success attained by LDA is due to the misclassification of three minor workers as C. quadrimaculatus. This is because the third variant in C. gibber and members of C. quadrimaculatus species share qualitative morphological traits, and both species display overlapping ranges of quantitative measurements. The grouping of C. gibber in the same cluster shown by the dendrogram of multivariate morphometric analysis corroborates the species hypothesized by the taxonomic revision based on qualitative morphology.
Camponotus egregius r. gouldi
Forel, 1886a: iv. Holotype (by monotypy) major worker, Madagascar (Grandidier) [not examined, type not found]. Combination in Camponotus (Myrmogigas): Forel, 1912: 91; in Camponotus (Dinomyrmex): Forel, 1914: 268; in Camponotus (Tanaemyrmex): Emery, 1925: 85. Raised to species:
Madagascar: Province Mahajanga: PN Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest, ground nest, 2–8 Apr 2001 (Fisher, Griswold & Malagasy Arthropod Team) collection code: BLF03536, specimen code: CASENT0437531 (
Madagascar: Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle (Rin’Ha, Mike) (
With head in full-face view, lateral margins of head anterior to eye level parallel and lacking erect hairs, posterior portion of head extending into a neck and anteromedian clypeal margin continuously forming broad convexity; dorsal outline of mesosoma broadly convex; body color black to dark brown.
Minor worker. In full-face view, head sides anterior to level of eye approximately parallel; lateral cephalic margins posterior to level of eye converging posteriorly and projecting into short neck; eye medium (EL/CS: 0.24±0.01; 0.23–0.26), protruding, and breaking lateral cephalic border, level of posterior margin located approximately at posterior 1/3 of head (PoOc/CL: 0.29±0.02; 0.26–0.32); frontal carinae more or less wide posteriorly (FR/CS: 0.26±0.01; 0.25–0.27), the same width as its smallest distance to eye; clypeus without anterolateral angle, anteromedian margin broadly rounded, mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.56±0.06; 1.44–1.66). Mesosoma presenting even convexity; promesonotum strongly convex, posterior portion of mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost flat, its junction to declivity bluntly angulate; propodeal dorsum 3 × as long as declivity. Petiole nodiform, with dorsal margin inclined posteriorly, forming a blunt angle to anterior margin, medially excised at the junction to posterior margin, anterior face 1/2 height of posterior face, which is inclined anteriorly to the dorsum; femur of hind leg rounded axially, not twisted basally.
Camponotus gouldi (neotype specimen) A lateral view B head in full-face view C dorsal view of major worker CASENT0437531.
First and second gastral tergites without a pair of white spots; lateral margin of head without erect hairs; erect hairs lacking near the end of neck; antennal scape without suberect hairs and covered with short appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum without erect hairs.
Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 5.19±0.23; 4.89–5.51; CWb/CL: 0.98±0.01; 0.97–1.00), which has no prolonged neck; the more strongly built mandible; antennal scape not extending beyond posterior cephalic margin; promesonotum convex; metanotal groove visible; propodeal dorsum feebly sloping towards declivity, its length the same as the height of declivity; petiolar node compressed anteroposteriorly.
The dry forest of the west, the spiny forest and thicket of the southwest, the transitional humid forest in the PN Andohahela, the savannah shrubland and woodland, and the Uapaca woodland and montane rainforest of the south-central high plateau of Madagascar are all habitats where C. gouldi occurs (Fig.
Camponotus gouldi A lateral view B head in full-face view C dorsal view of minor worker CASENT0121617 D distribution map.
Camponotus gouldi is mostly similar to C. aro, but the posterior portion of the head for the latter is normally rounded, not extending into a short neck, and the propodeal dorsum immediately posterior to the metanotal groove is convex, then becomes concave medially and rounds to the declivity surface.
For C. gouldi, the decisions drawn from qualitative morphology-based taxonomy agree with the classification hypothesis provided by the exploratory data analysis and the cumulative LDA of the multivariate morphometrics. The combination of this information corroborates the status of C. gouldi as a species.
Neotype designation has been made for C. gouldi because its type is presumed lost. No type specimen could be found at the renowned museums in Europe that might have held the specimen. One of the author BLF visited the Forel collection at
Camponotus rubripes r. hagensii
Forel, 1886a: 158. Lectotype minor worker, by present designation, Centre de Madagascar (Hildebrandt) AntWeb CASENT0910112 (
Madagascar: Antananarivo: Central Madagascar; Ambatomanjaka; Miarinarivo, -18.876091, 46.865775, 1343 m (Hildebrandt) (
In full-face view, lateral margin of head anterior to eye level diverging posteriorly and covered with erect hairs; anterior clypeal margin truncate; mesosoma in profile low and long; gastral tergites without abundant pubescence; head and gaster black, mesosoma reddish orange to brown.
Minor worker. In full-face view, head sides diverging towards broadly convex posterior margin or parallel anterior to eye level and rounding evenly to posterior border; eye protruding and large (EL/CS: 0.30±0.01; 0.28–0.33), breaking lateral cephalic margin; level of its posterior margin located ca. at posterior 1/4 of head (PoOc/CL: 0.25±0.01; 0.23–0.25); frontal carinae wide (FR/CS: 0.34±0.01; 0.31–0.35), posteriorly diverging, distance between them larger than their smallest distance to eye; clypeus without well-defined anterolateral corner, anteromedian margin approximately truncate; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.24±0.06; 1.09–1.36). Promesonotum weakly convex, mesonotum with posterior portion flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight, its junction to declivity bluntly angulate; height of propodeal declivity 3/4 length of dorsum. Petiolar node short and high; its dorsal margin inclined posteriorly, rounding to anterior margin; height of anterior face 2/3 that of posterior; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head lacking; more than six erect hairs present near posterior margin of head; antennal scape covered only with suberect hairs inclined at ca. 30° and appressed hairs; pronotum with few erect hairs, mesonotum bearing a pair of erect hairs; posterodorsal corner of propodeum with two pairs of erect hairs. Head, antenna, and gaster dark brown or reddish brown; mesosoma, petiolar node, and legs yellow to orange-yellow.
Major worker. Differing from minor worker in having larger, heart-shaped head (CS: 2.37±0.26; 2.00–2.70; CWb/CL: 0.97±0.05; 0.92–1.04) with broadly concave posterior margin, anteromedian margin of clypeus slightly excised medially; apical 1/4 of antennal scape extending beyond posterior cephalic margin; robust mesosoma with distinct metanotum; propodeal dorsum convex immediately behind metanotum, < 2 × the height of declivity surface; petiolar node much higher than long and tapering dorsally.
Camponotus hagensii is known mostly from montane rainforest, montane ericoid thicket, philipia forest, montane shrubland, and savannah grassland of the high plateau ranging from the PN Marojejy in the north to the Anosyenne Mountains in the south (Fig.
Camponotus hagensii A lateral view B head in full-face view C dorsal view of minor worker CASENT0191568 D distribution map.
Camponotus hagensii is similar to C. aurosus with respect to the bicolored body and the presence of erect hairs on the lateral margin of the head anterior to eye level, but in C. aurosus the anterior clypeal margin is broadly triangular, the mesosoma is short and high in profile, and the gastral tergites are covered with abundant pubescence.
The cluster of C. hagensii samples presented by the dendrogram of multivariate morphometric analysis is supported by the cumulative LDA at 100% identification success. This finding is consistent with the results of the qualitative morphology-based study.
Madagascar: Province Antsiranana: Forêt d’Analabe, 30 km 72° ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest, ground forager, 28 Nov 2003 (B.L. Fisher et al.) collection code: BLF09544, specimen code: CASENT0499207 (
1 minor worker of same data as holotype but with collection code: BLF09463 and specimen code: CASENT0499217 (
Madagascar: Antsiranana: Forêt d’Analabe, 30 km 72° ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher) (
With head in full-face view, eye not breaking lateral cephalic margin; mesonotum short and lacking constriction; propodeal dorsum transversely medially concave at ca. posterior 1/2; petiolar node ca. as high as long.
Minor worker. With head in full-face view, lateral margins anterior to eye level approximately parallel, progressively rounding evenly towards slightly concave rear margin; eye protruding and large (EL/CS: 0.25±0.01; 0.25–0.26), not breaking lateral cephalic margin, location of its posterior margin at posterior 1/3 of head (PoOc/CL: 0.28±0.01; 0.27–0.29); frontal carinae not widely opened posteriorly, (FR/CS: 0.25±0.00; 0.24–0.25); clypeus without anterolateral angle, anteromedian margin broadly convex; mandible with two apical teeth normally spaced; antennal scape relatively long (SL/CS: 1.63±0.05; 1.58–1.70). Promesonotum noticeably convex, metanotal groove noticeably visible; anterior portion of propodeal dorsum convex, then concave and becoming straight with blunt angle to declivity surface. Petiole nodiform, anterior and posterior margins approximately the same height, dorsal margin separated by an angle from the anterior and gently rounding to the posterior.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head present anterior to and behind eye level; posterior margin of head with two elongate, erect hairs; antennal scape covered with erect hairs; junction of propodeal dorsum and declivity with two pairs of erect hairs. Body color orange-brown with darker coxa, junction of segments, abdominal sternites, and appendices.
Unknown.
Endemic to Madagascar, C. harenarum is only known from the littoral rainforest of Analabe in Daraina, in the northeastern region of the island (Fig.
Camponotus harenarum A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0499207 D distribution map.
Camponotus harenarum can be confounded with C. karsti, but the latter has a more sloping anterior portion of the propodeum that makes a noticeable angle with the posterior region, and a shorter anterior margin of the petiolar node.
The species name harenarum is a Latin plural noun in the genitive case of harena, which means sand. It is in reference to the littoral rainforest, the only habitat in which it has been found.
Camponotus maculatus r. hova
Forel, 1891: 35. Syntype workers and queen, Morondava côte ouest de Madagascar (Grandidier) (
Camponotus hova var. obscuratus
Emery, 1925: 85. Syntype workers and male, SW Madagascar (Voeltzkow) (
Europa Island: Europa Island (Voeltzkow) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible closely spaced; antennal scape covered with erect hairs.
Minor worker. With head in full-face view, lateral margins of head anterior to level of eye parallel and converging progressively to posterior margin; head sides behind eye level ca. 1/4 length of head (PoOc/CL: 0.25±0.02; 0.22–0.28). Eyes protruding and large (EL/CS: 0.30±0.01; 0.28–0.32), breaking lateral cephalic margins; frontal carinae not widely diverging posteriorly (FR/CS: 0.27±0.01; 0.24–0.30), posteriorly parallel; clypeus with anterolateral angle and triangular or convex anteromedian margin; two apical teeth of mandible closely spaced; antennal scape relatively long (SL/CS: 1.53±0.08; 1.41–1.83). Dorsum of mesosoma highly convex, mesonotum with posterior portion flat immediately anterior to metanotal groove; metanotal groove weakly visible; propodeal dorsum almost straight, junction to declivity with blunt angle; declivity height 1/2 length of propodeal dorsum; petiolar node nodiform with dorsal margin inclined posteriorly and forming a blunt angle to anterior face; anterior face of petiolar node 1/3 height of posterior face. Tibia of hind leg rounded axially and not twisted basally.
First and second gastral tergites without a pair of white spots; lateral margin of head with erect hairs; posterior margin of head with more than six erect hairs. Antennal scape covered with suberect hairs inclined ca. 30° and abundant pubescence. Posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 3.17±0.16; 2.94–3.48; CWb/CL: 0.94±0.03; 0.89–1.01) with broadly concave posterior margin; two apical teeth of mandible normally spaced; apical 1/4 of antennal scape surpassing posterior cephalic margin; robust mesosoma, metanotum distinctly visible, propodeal dorsum joining declivity in broad angle; dorsal margin of petiolar node inclined posteriorly from shorter anterior face towards much longer posterior face. More pairs of erect hairs on promesonotum, junction of propodeal dorsum and declivity, and posterodorsal margin of petiolar node.
Camponotus hova is a widespread species that occurs in dry forests of the west from the north throughout the center and the southwest of Madagascar, and is also known from Juan de Nova, Europa, and Mayotte islands (Fig.
Camponotus hova A lateral view B head in full-face view C dorsal view of minor worker CASENT0055710 D distribution map.
Camponotus hova shows significant morphological variation across its wide distribution in Madagascar and nearby islands. Two variants are recognized according to the forms of the dorsum of the propodeum, but these merge gradually into the typical form through their geographical distribution.
Variant 1. Workers of this variant express the typical form by which the posterior 1/2 of the mesonotum to the posterodorsal corner of the propodeum is straight in profile. The petiolar node is characterized by the posterior inclination of the dorsal margin toward the anterior face, which is ca. 1/3 the height of the posterior face. This variant occupies the southwest region of Madagascar and surrounding islands. Integument is brown to dark brown or blackish brown in color.
Variant 2. This variant is known from the northwestern part of Madagascar and is characterized by the presence of a slightly broad concavity from the posterior 1/2 of mesonotum to posterodorsal corner of propodeum in lateral view and by the pale yellow to reddish orange color the body. Its petiole is nodiform, and the dorsal margin rounds to the anterior face, which is 1/2 height of the posterior face.
Based on the original description of C. hova (Forel, 1891) and C. hova obscuratus (Emery, 1925) and the examination of their respective syntype specimens, there are no distinctive characters that were found to differentiate both taxa. The observation of the collection obtained from the recent survey of the Malagasy ant fauna indicates that the distinctive characters of C. hova obscuratus vary within and across the populations of C. hova. Therefore, it is reasonable to place safely C. hova obscuratus in synonymy here.
Although qualitative morphology recognized the two variants in C. hova, the exploratory analysis of NC-clustering did not completely detect their distinction. The combination of the two variants is classified by confirmatory LDA at 100% of success.
Camponotus hovahovoides
Forel, 1892: 232. Syntype minor and major workers, queen and male, Madagascar, Andrangoloaka (Sikora) (
Camponotus radamae var. hovoides
Dalla Torre, 1893: 249. Syntype minor and major workers, queen and male, Madagascar; 1 syntype minor worker designated as lectotype, by present designation, Antananarivo (Camboué) AntWeb CASENT0101421 (
Madagascar: Antananarivo: 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (Fisher, Griswold et al.) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible normally spaced; antennal scape covered with suberect hairs; in lateral view, dorsum of mesosoma from mid-mesonotum to posterodorsal corner of propodeum approximately straight, propodeal dorsum ca. 3 × as long as the height of declivity surface; petiolar node flattened anteroposteriorly.
Minor worker. Head sides anterior to level of eye parallel; lateral margins posterior to level of eye converging progressively to posterior margin; length of posterior portion of head behind eye level 1/3 length of head (PoOc/CL: 0.28±0.01; 0.25–0.31). Eyes protruding and large (EL/CS: 0.30±0.01; 0.27–0.34), breaking lateral cephalic margin. Frontal carinae strongly diverging posteriorly (FR/CS: 0.30±0.01; 0.28–0.33); clypeus projecting into anterolateral angle and medially with blunt angle or convex. Two apical teeth of mandible distantly spaced. Antennal scape relatively long (SL/CS: 1.54±0.08; 1.43–1.75) with erect to suberect hairs inclined to 45°. Promesonotum weakly convex and mesopropodeum almost flat, metanotal groove weakly visible, propodeal dorsum anteriorly convex and posteriorly flat, joining the declivity in a blunt angle, propodeal declivity height 1/2 dorsum length. Petiolar node flattened anteroposteriorly or short and high, its dorsal margin rounding to anterior margin. Tibia of hind leg rounded and not twisted dorsoventrally.
First and second gastral tergites without a pair of white spots. Lateral margin of head covered with erect hairs; posterior margin of head with two or three pairs of erect hairs; posterodorsal angle of propodeum with one or two pairs of erect hairs.
Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 2.66±0.20; 2.40–3.07; CWb/CL: 0.77±0.02; 0.72–0.80); the more strongly built mandible; apical 1/5 of antennal scape surpassing posterior cephalic margin; metanotum visible; propodeal dorsum convex and its junction to declivity broadly angulate; petiolar node much higher than long.
A widely distributed species, C. hovahovoides occurs mainly in the eastern lowland to montane rainforests and montane shrublands of Madagascar (Fig.
Camponotus hovahovoides A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT0496908 and CASENT0487242 G distribution map.
See also discussion under C. mixtellus. Camponotus hovahovoides is one of the more common species of Myrmosaga occupying the montane rainforests of Madagascar and shows notable morphological variation. Two variants are recognized based on the shapes of the mesosoma and body colors, but these merge progressively into one typical form across the geographic distribution of the species.
Variant 1. In this variant the posterior 1/2 of mesonotum to posterodorsal corner of propodeum is not straight in lateral view and the body is pale yellow to reddish orange in color.
Variant 2. This variant is known by a dorsum of mesosoma from mid-mesonotum to posterodorsal corner of propodeum that is approximately straight, and its body is dark brown to black in color or the mesosoma is lighter in color relative to the darker head and gaster. Workers of this variant have been collected from mountaintops.
Camponotus hovahovoides has a wide distribution across the humid forest in Madagascar, this information combined with the results of the comparison of the syntype specimens of C. radamae var. hovoides with those of C. hovahovoides indicate that there is no strong morphological difference between both taxa. The few specimens of C. radamae var. hovoides represent a rare variation of C. hovahovoides across its geographic distribution.
The NC-Clustering technique was used to reveal the two variants and demonstrates that the quantitative morphological characters did not identify the difference between both variants. The members of each variant are scattered along the cluster of C. hovahovoides. More robust molecular phylogenetic information is needed to determine whether the variants constitute separate species and to study the ecological and evolutionary forces underlying these morphological variations.
With head in full-face view, posterior portion of head extending into a neck, and anteromedian clypeal margin continuously forming broad convexity; dorsal outline of mesosoma complex; body color black to dark brown.
See the geographic distribution and biology in
Camponotus imitator can be easily separated from the other species of the Myrmosaga group in that it is the only species that is characterized by an elongate mesonotum that is constricted at midlength, and its propodeum is broadly convex when the mesosoma is viewed in profile.
See also discussion under the section “Morphological diagnosis of the worker caste of the Malagasy Myrmosaga”.
The grouping of C. imitator revealed by the NC-clustering method dendrogram is confirmed by the cumulative LDA at 100% identification success, indicating the support of the species hypothesized by qualitative morphology-based taxonomy.
Camponotus quadrimaculatus var. immaculatus
Forel, 1892: 233. Syntype workers, Madagascar Andrangoloaka (Sikora); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102430 (
Camponotus quadrimaculatus opacata
Emery, 1925: 123. Syntype workers and alate queen, Madagascar, Baie d’Antongil (Mocquerys); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102107) (
Madagascar: Antananarivo: [Andrangoloaka]; Mantasoa; Manjakandriana, -19.033333, 47.9166666, 1409 m (Sikora) (
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; white spots absent on second and third abdominal tergite.
Minor worker. With head in full-face view, lateral margins diverging towards almost straight posterior margin; eye slightly convex and small (EL/CS: 0.25±0.01; 0.23–0.28), not interrupting lateral cephalic border, level of its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.26±0.02; 0.22–0.28); frontal carinae wide, posteriorly diverging (FR/CS: 0.32±0.01; 0.30–0.33), distance between them larger than their smallest distance to eye; clypeus with bluntly angulate anterolateral corner and straight anteromedian margin; two apical teeth of mandible normally distant; antennal scape relatively short (SL/CS: 1.28±0.05; 1.19–1.34). Promesonotum evenly convex; posterior portion of mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum strongly concave medially; junction of dorsal margin of propodeum with declivity bluntly angulate; propodeal dorsum ca. 2 × as long as declivity. Petiole nodiform, its dorsal margin inclined posteriorly and rounding to anterior margin; anterior and posterior faces almost the same height; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head absent; two erect hairs present near posterior cephalic margin; antennal scape with appressed hairs only; pronotum and mesonotum with a pair of erect hairs each, posterodorsal angle of propodeum with a pair of erect hairs. Integument shining; body color dark brown; distal portion of leg reddish brown.
Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 2.32±0.12; 2.13–2.46; CWb/CL: 1.00±0.03; 0.96–1.04), with slight concavity of posterior margin; anteromedian margin of clypeus noticeably concave; apical 1/4 of antennal scape extending beyond posterior cephalic margin; robust mesosoma, promesonotum convex, propodeal dorsum straight immediately posterior to metanotum, feebly concave towards declivity, length the same as height of declivity; petiolar node tapering dorsally.
Occupying the high plateau of Madagascar, C. immaculatus occurs in shrubland and grassland areas, montane rainforest, savannah grassland, and Uapaca woodland habitats (Fig.
Camponotus immaculatus A lateral view B head in full-face view C dorsal view of minor worker CASENT0082438 D distribution map.
Camponotus immaculatus can be easily separated from C. kelimaso and C. lubbocki by the presence of the broad concavity on the propodeal dorsum. It also can be distinguished from other species because the anteromedian margin of its clypeus is truncate and no white spots are visible on the abdominal tergites.
When describing C. quadrimaculatus opacata,
The delimitation argument for C. immaculatus is strengthened by the congruence between the results of traditional qualitative morphology and the NC-clustering method. However, LDA produces a high 8.4% error rate in the classification of C. immaculatus, due to a misidentification of a single minor worker as C. quadrimaculatus. The high error rate is due in large part to the small range of minor worker forms of C. immaculatus available for this study.
Madagascar: Province Antsiranana: Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest, 11 Apr 2009 (Fisher, Griswold et al.) collection code: BLF03132, specimen code: CASENT0408908 (
2 minor workers with same data as holotype but respectively specimen coded as: CASENT0408907, CASENT0408909 (
Madagascar: Antsiranana: Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest (Fisher, Griswold et al.) (
In full-face view, lateral margins of head anterior to eye level approximately parallel and covered with erect hairs; two apical teeth of mandible normally spaced; mesonotum short and lacking constriction; propodeal dorsum approximately straight, 2 × as long as declivity; dorsal margin of petiole shorter than posterior margin.
Minor worker. In full-face view, lateral margins of head anterior to eye level approximately parallel, rounding progressively towards a slightly concave posterior margin; eye convex and large (EL/CS: 0.27±0.00; 0.27–0.28), not breaking lateral cephalic margin, location of its posterior margin at posterior 1/4 of head (PoOc/CL: 0.26±0.00); frontal carinae parallel, not widely opened posteriorly (FR/CS: 0.27±0.0); clypeus lacking anterolateral angle, anteromedian margin broadly convex; two apical teeth of mandible normally spaced; antennal scape relatively long (SL/CS: 1.62±0.01; 1.61–1.63). Dorsal outline of mesosoma approximately evenly convex; metanotal groove visible; propodeal dorsum straight, ca. 2 × as long as declivity, joining it at a blunt angle. Petiole nodiform, its dorsal margin shorter than anteriorly inclined rear margin and joining the anterior margin at a blunt angle.
First and second gastral tergites without a pair of white spots; short erect hairs on lateral margin of head present; near posterior margin of head with two elongate erect hairs; antennal scape covered with erect hairs; junction of propodeal dorsum and declivity with one pair of erect hairs.
Major worker. Unknown.
Camponotus joany is geographically restricted to the dry forest of Montagne de Francais in the north of Madagascar (Fig.
Camponotus joany A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0408908 D distribution map.
See discussion under C. aina.
The species name joany is a non-Latin singular noun used in apposition.
Madagascar: Province Antsiranana: RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, tropical dry forest, low vegetation, 27 Nov 1990 (P.S. Ward) collection code: PSW11005, specimen code: CASENT0217292 (
1 minor worker with same data as holotype but respectively specimen coded as: CASENT0837940 (
Madagascar: Antsiranana: RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, Dry Forest, (P.S. Ward) (
With head in full-face view, eye not breaking lateral cephalic margin; mesonotum short and lacking constriction; promesonotum an even convexity; propodeum with blunt angle at ca. posterior 1/2; anterior margin of petiolar node in profile very low, dorsal surface noticeably horizontal, ca. 2 × as long as posterior surface.
Minor worker. With head in full-face view, lateral margins anterior to eye level approximately parallel, progressively rounding evenly towards slightly concave rear margin; eye protruding and large (EL/CL: 0.26±0.01; 0.25–0.26), not breaking lateral cephalic margin, location of its posterior margin at posterior 1/4 of head (PoOc/CL: 0.26±0.01; 0.25–0.27); frontal carinae parallel, not widely opened posteriorly (FR/CS: 0.25); clypeus with blunt or poorly defined anterolateral angle, anteromedian margin broadly convex; two apical teeth of mandible normally spaced; antennal scape relatively long (SL/CS: 1.65±0.03; 1.63–1.67). Promesonotum noticeably convex, followed by a slightly visible metanotal groove; anterior portion of propodeal dorsum sloping along its first 1/2 then slightly convex and rounding to declivity surface. Anterior margin of petiolar node ca. 1/2 height of posterior margin, dorsal margin separated by an angle from anterior margin and inclined posteriorly to jointly rounding to posterior margin.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head present on anterior and behind eye level; posterior margin of head with two elongate, erect hairs; antennal scape covered with appressed hairs; junction of propodeal dorsum and declivity with two pairs of erect hairs.
Major worker. Unknown.
The distribution of C. karsti is limited to the dry forest of the RS Ankarana in the north of Madagascar (Fig.
Camponotus karsti A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217292 D distribution map.
Camponotus karsti can be confounded with C. harenarum because both species are characterized by the presence of a blunt angle at approximately the posterior 1/2 of propodeal dorsum. However, C. harenarum can be distinguished by the separate convexity of its mesonotum and anterior 1/2 of propodeum, and its petiolar node is approximately as high as long.
The species name karsti is a non-Latin singular noun used in apposition.
Madagascar: Province Antsiranana: PN Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, -14.435, 49.76, 775 m, rainforest, ex rotten log, 16 Nov 2003 (B.L. Fisher et al.) collection code: BLF09010, specimen code: CASENT0487718 (
3 minor workers of same data as holotype but specimens coded as: CASENT0837637, CASENT0837636, CASENT0837635 (
Madagascar: Antsiranana: Makirovana forest, -14.16666, 49.95, 715 m, rainforest (B.L. Fisher et al.) (
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; dorsum of second and third abdominal tergites lacking white spot; propodeal dorsum straight; body color dark brown; antennal scape without erect hairs.
Minor worker. In full-face view, head sides diverging towards anterior to level of eye, evenly rounding to approximately straight posterior margin; head widest at eye level; eyes almost flattened and small (EL/CS: 0.19±0.01; 0.16–0.21), not breaking lateral cephalic margin, level of posterior margin located approximately at posterior 1/3 of head (PoOc/CL: 0.28±0.01; 0.27–0.29); frontal carinae more or less wide (FR/CS: 0.31±0.02; 0.28–0.34), posteriorly diverging, distance between them smaller than their smallest distance to eye; clypeus with anterolateral angle and approximately straight anteromedian margin; mandible with two apical teeth distantly spaced; antennal scape relatively short (SL/CS: 1.19±0.05; 1.11–1.27). Promesonotum weakly convex; mesonotum with posterior portion flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight, its angle to declivity widely rounded; propodeal dorsum ca. 2 × as long as declivity. Petiolar node flattened anteroposteriorly, without obvious dorsal margin; anterior face almost the same height as posterior face; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; lateral cephalic margin with erect hairs anterior to level of eyes; no erect hairs on lateral margin of head posterior to eye level; two erect hairs present near posterior margin; antennal scape only covered with appressed hairs; pronotum with few erect hairs; mesonotum with a pair of erect hairs; two erect hairs present on anterior to posterodorsal corner of propodeum. Body color reddish brown to pale brown.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 2.83±0.25; 2.35–3.15; CWb/CL: 0.99±0.04; 0.92–1.04) with broadly concave posterior margin; apical 1/4 of antennal scape surpassing posterior cephalic margin; robust mesosoma with separate convexity of promesonotum, lower position of metanotum and propodeum, propodeal dorsum approximately the same length as declivity surface and with rounded junction; petiolar node more flattened anteroposteriorly.
The distribution of C. kelimaso is limited to the eastern lowland rainforest of Madagascar between the Makirovana forest in the north and the Forêt d’Ambalagoavy Nord in the south (Fig.
Camponotus kelimaso A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0487718 D distribution map.
Camponotus kelimaso can be separated from C. immaculatus by its straight propodeal dorsum. It can be differentiated from C. lubbocki by the approximately straight posterior margin of the head and its small compound eyes.
Based on the information provided by the NC-clustering method, the grouping of C. kelimaso supports the distinction of the species by conventional qualitative taxonomy. The confirmatory LDA identified the samples successfully at 100%.
The species name kelimaso is a non-latin word derived from the Malagasy word for “small eye”. It refers to the fact that its compound eyes are small compared to those of other species in the subgenus Myrmosaga.
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin broadly triangular.
In the present study, more samples have been attributed to C. liandia. The species is generally known from coastal scrub and littoral forest in the east, montane rainforest, grassland, shrubland, savannah grassland, woodland, and Uapaca woodland on the central high plateau (Fig.
Camponotus liandia A lateral view B head in full-face view C dorsal view of minor worker CASENT0491145 D distribution map.
With the additional samples, C. liandia presents two morphological variations: a matte integument covered with microreticulate sculpture, and a smooth and shining body. The first form has been observed from samples collected from the Forêt d’Atsirakambiaty in the grassland of the south-central high plateau. The second form occupies coastal scrub and littoral forest in the east and dry forest on Tsingy and burned savannah in the west.
Camponotus liandia was originally described under the subgenus Mayria (
The cluster for C. liandia is composed of a larger number of worker samples to include the qualitative morphological variation within the species. The results of the exploratory analysis using NC-clustering methods are congruent with the definition of C. liandia based on traditional qualitative morphology. Identification of this species is confirmed by LDA with 100% success.
Madagascar: Province Antsiranana: RNI Lokobe, 6.3 km 112° ESE Hellville, Nosy Be, -13.41933, 48.33117, 30 m, rainforest, ex rotten log, 19–24 Feb 2001, (Fisher, Griswold et al.) collection code: BLF03476, specimen code: CASENT0436584 (
2 workers of same data as holotype but specimen coded as: CASENT0835595 (
Madagascar: Antsiranana: Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (
With head in full-face view, lateral cephalic margins converging posteriorly towards eye level, then rounding to occipital corner of head; anteromedian margin of clypeus noticeably excised medially; two apical teeth of mandible normally spaced; lateral cephalic margin anterior to eye level covered with erect hairs.
Minor worker. With head in full-face view, lateral cephalic borders anterior to level of eye parallel to each other, converging strongly towards posterior margin behind eye level; eye large and convex (EL/CS: 0.30±0.01; 0.27–0.31), breaking lateral margin of head, level of its posterior margin located approximately at posterior 1/3 of head (PoOc/CL: 0.29±0.01; 0.28–0.31); frontal carinae not widely opened (FR/CS: 0.26±0.01;0.25–0.27), posteriorly parallel, distance between them larger than smallest distance to eye; clypeus with anterolateral angle and medially slightly concave anteromedian margin; two apical teeth of mandible distantly spaced; antennal scape relatively long (SL/CS: 1.80±0.03; 1.76–1.85). Mesosoma long and low (MPH/ML: 0.29±0.01; 0.29–0.30), with its dorsal outline almost flat; metanotal groove feebly visible; propodeal dorsum feebly convex anteriorly and flat posteriorly; junction of propodeal dorsum to declivity bluntly angulate; propodeal declivity 1/3 length of the dorsum. Petiole nodiform, tapering dorsally; its dorsal margin inclined posteriorly and forming a blunt angle to anterior face; posterior face 2 × as high as the anterior; femur of hind leg rounded axially, without twist near base.
First and second gastral tergites without a pair of white spots; with head in full-face view lateral margin covered with erect hairs; near posterior margin of head with a pair of erect hairs; antennal scape covered with erect hairs inclined at ca. 45°; pronotum with numerous erect to suberect hairs; a pair of erect hairs present on posterodorsal angle of propodeum.
Major worker. Unknown.
Camponotus lokobe is endemic to Madagascar and restricted to the rainforest of the RNI Lokobe in the north of the island (Fig.
Camponotus lokobe A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0436584 D distribution map.
Camponotus lokobe can be separated from C. dufouri and C. tendryi by the fact that its clypeus is characterized by a medially excised anteromedian margin.
The grouping of C. lokobe in the same cluster shown by the dendrogram of multivariate morphometric analysis is confirmed by the cumulative LDA at 100% identification success, corroborating the species hypothesized by the taxonomic revision based on qualitative morphology.
The species name lokobe is a singular non-Latin noun used in apposition and refers to the type locality.
In full-face view, lateral margin of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; no white spot on dorsum of second and third abdominal tergites; body color black; antennal scape without erect hairs, propodeal dorsum slightly concave.
See the geographic distribution and biology in
Camponotus lubbocki might be confused with C. liandia but the latter has a broadly convex anteromedian margin of the clypeus. It looks similar to C. immaculatus and C. kelimaso because they too lack white spots on the second and third abdominal segments, but in C. immaculatus the propodeal dorsum is transversely concave and in C. kelimaso the eyes are small and the posterior cephalic margin is approximately straight.
The species delimitation of C. lubbocki based on qualitative morphology is sustained by the NC-clustering method. Identification of this species is confirmed by LDA with 100% success (see also
Madagascar: Province Toliara: Androy Region, District of Tsihombe, 74 km S of Tsihombe, Reserve Speciale Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush, Malaise trap, 30 Apr–11 May 2003, (Rin’ha, Mike) CASENT0115206 (
2 minor workers of same data as holotype but respectively with collection codes: MG-23-34 and MG-23-33 and specimen codes: CASENT0115287 and CASENT0115718 (
Madagascar: Toliara: Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush, (Mike, Frank Parker, Rin’ha) (
With head in full-face view, lateral margins of head anterior to eye level parallel and lacking erect hairs, lateral and anteromedian clypeal margins continuously forming broad convexity; scape without erect hairs; mesosoma long and low; body color yellowish to brown.
Minor worker. In full-face view, head sides anterior to level of eye parallel, rounding evenly to posterior margin behind eye level; eye protruding and large (EL/CS: 0.32±0.01; 0.30–0.35), breaking lateral cephalic margin, level of its posterior margin situated approximately at posterior 1/4 of head (PoOc/CL: 0.23±0.01; 0.20–0.24); frontal carinae close to each other (FR/CS: 0.27±0.01; 0.25–0.28), posteriorly parallel, distance between them smaller than their smallest distance to eye; clypeus without well-defined anterolateral angle, its anteromedian margin broadly convex; mandible with two apical teeth normally spaced; antennal scape relatively long (SL/CS: 1.40±0.04; 1.33–1.45). Promesonotum weakly convex; mesonotum with posterior portion flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight (MPH/ML: 0.37±0.03; 0.34–0.50); rounding to declivity, 2 × as long as declivity. Petiolar node short and high, with dorsal margin inclined posteriorly, rounding to anterior margin; height of anterior face 2/3 of that of posterior face; femur of hind leg rounded axially, without twist near base.
First and second gastral tergites without a pair of white spots; lateral margin of head anterior to eye level, with no erect hairs posterior to eye level; two erect hairs present close to posterior margin; antennal scape covered only with appressed hairs; erect hairs lacking on promesonotum; posterodorsal corner of propodeum with a pair of erect hairs. Integument shining; body color yellowish.
Major worker. Unknown.
Camponotus mahafaly is geographically restricted to dry forest and spiny bush and forest habitats of the southwest and south of Madagascar (Fig.
Camponotus mahafaly A lateral view B head in full-face view C dorsal view of minor worker CASENT0115287 D distribution map.
Camponotus mahafaly might be confounded with C. cemeryi because the petiole of both species is not nodelike, but in the latter species the antennal scape is covered with suberect hairs and the mesosoma is generally short and high, with a dorsal outline that is strongly convex.
The qualitative morphology-based study of this species agrees with the multivariate morphometrics to support the taxonomic determination of C. mahafaly. Separation of this species is confirmed by LDA with 100% success.
The species name mahafaly is a singular non-Latin noun used in apposition and refers to the Mahafaly region in the southwest of Madagascar.
Camponotus radamae mixtellus
Dalla Torre, 1893: 249 Lectotype minor worker, by present designation, Madagascar, Forêt des bords de l’Ivondrona, près de Tamatave (Dr Conrad Keller) AntWeb CASENT0101893 (
Comoros: Anjouan: Mount Ntringui, -12.19865, 44.41866, 740 m, montane forest (B.L. Fisher et al.) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible normally spaced; antennal scape covered with suberect hairs inclined at ca. 30°; in lateral view, posterior 1/2 of mesonotum to posterodorsal corner of propodeum somewhat convex, propodeal dorsum ca. 2 × as long as the height of declivity surface; petiolar node flattened anteroposteriorly.
Minor worker. In full-face view, head sides anterior to level of eye parallel; lateral margins posterior to level of eye converging progressively to posterior margin; length of posterior portion of head behind eye level 1/3 length of head (PoOc/CL: 0.30±0.01; 0.28–0.31). Eyes protruding and large (EL/CS: 0.30±0.02; 0.27–0.33), breaking lateral cephalic margin. Frontal carinae not strongly diverging posteriorly (FR/CS: 0.28±0.01; 0.26–0.30) and posteriorly parallel; clypeus with anterolateral angle and its anterior margin with medial blunt angle or convex. Mandible with six teeth, two apical teeth normally placed. Antennal scape relatively long (SL/CS: 1.55±0.07; 1.40–1.73) with suberect hairs inclined 30°. Promesonotum weakly convex and mesopropodeum feebly convex (MPH/ML: 0.39±0.02; 0.36–0.42), posterior portion of mesonotum flat immediately anterior to metanotal groove, metanotal groove weakly visible, propodeal dorsum anteriorly convex and posteriorly flat, junction of dorsal margin and declivity with blunt angle, propodeal dorsum 2 × as long as the declivity. Petiolar node flattened anteroposteriorly or short and high, its dorsal margin rounding to anterior margin. Tibia of hind leg rounded.
First and second gastral tergites without a pair of white spots. Erect hairs on lateral margin of head anterior to level of eyes present, but absent posterior to level of eyes; posterior margin of head with two pairs of erect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 2.92±0.15; 2.62–3.10; CWb/CL: 0.93±0.03; 0.90–0.99), in full-face view with slightly concave posterior margin, eye not breaking lateral cephalic margin; apical 1/4 of antennal scape surpassing posterior cephalic margin; robust mandibles and mesosoma, metanotum distinctly visible; petiolar node tapering dorsally.
Camponotus mixtellus is a widespread species of the Malagasy region that occurs in the eastern lowland to montane rainforests and part of the western dry forests of Madagascar; it is known also from secondary rainforest and dry forest habitats of Grand Comore and Moheli of the Comoros islands (Fig.
Camponotus mixtellus A lateral view B head in full-face view C dorsal view of minor worker CASENT0076675 D distribution map.
Camponotus mixtellus can be confused with C. hovahovoides, but the latter has a longer propodeal dorsum that is ca. 3 × as long as the declivity. It looks similar to C. boivini but the latter is characterized by an antennal scape covered with suberect hairs inclined at ca. 45°.
The cluster of C. mixtellus shown by the NC-clustering method is confirmed by the cumulative LDA at 100% identification success, indicating support of the species hypothesized by qualitative morphology-based taxonomy.
Madagascar: Province Antsiranana: PN Montagne d’Ambre, Antomboka, -12.50035, 49.175, 885 m, montane rainforest, ex rotten log, 16 Nov 2007 (B.L. Fisher et al.) collection code: BLF18356 specimen code: CASENT0134004 (
1 major worker of same data as holotype but with specimen code: CASENT0134003 (
Madagascar: Antsiranana: Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (
With head in full-face view, lateral cephalic margin converging posteriorly towards eye level, covered with erect hairs anterior and posterior to eye level; anteromedian margin of clypeus broadly convex; two apical teeth of mandible normally spaced.
Minor worker. In full-face view, head widest at midlength, lateral margins posterior to level of eye rounding evenly to posterior margin; eye large, slightly convex (EL/CS: 0.27±0.01; 0.26–0.30), not breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/4 of the head (PoOc/CL: 0.28±0.02; 0.25–0.32); frontal carinae wide (FR/CS: 0.25±0.01; 0.24–0.28), distance between them larger than their distance to eye; clypeus with anterolateral angle and anteromedian margin with blunt or convex angle; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.70±0.11; 1.59–1.88). Promesonotum weakly convex, posterior portion of mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight; dorsal margin of propodeum and declivity junction bluntly rounded; propodeal dorsum 2 × as long as declivity. Petiolar node short and high, its dorsal margin inclined posteriorly and joining anterior face in a blunt angle; anterior face 1/3 of height of the posterior; femur of hind leg rounded axially, without torsion near base.
First and second gastral tergites without a pair of white spots; erect hairs present on lateral margin of head, near its posterior margin bearing more than six erect hairs; antennal scape with suberect hairs inclined at ca. 30°, appressed hairs lacking; promesonotum covered with erect hairs; posterodorsal angle of propodeum with more than two erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 4.02±0.35; 3.69–4.61; CWb/CL: 0.92±0.08; 0.79–1.01); more robust mandible; apical 1/4 of antennal scape surpassing posterior cephalic margin; pronotum convex, mesonotum sloping towards metanotum, propodeal dorsum feebly convex and joining the declivity at a broad angle; petiolar node higher than long.
Camponotus niavo is geographically restricted to the transitional humid forest of Ampasindava Peninsula, the rainforests of the Galoko chain and RS Manongarivo, and the montane rainforest of the PN Montagne d’Ambre (Fig.
Camponotus niavo A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0134004 D distribution map.
See discussion under C. cervicalis.
The qualitative, morphology-based analysis of this species agrees with the multivariate morphometric analysis to provide 100% support for the taxonomic determination of C. niavo.
The species name niavo is a non-Latin singular noun used in apposition and is derived from the Malagasy word that means “from above”. It refers to the high mountains where the species has been collected.
Camponotus quadrimaculatus
Forel, 1886a: cii. Lectotype minor worker, by present designation, Madagascar, Fianarantsoa (Besson) AntWeb CASENT0102426 (
Camponotus quadrimaculatus sellaris
Emery, 1895: 344. Syntype workers, Madagascar, Antsiranana [[Diego Suarez; 7]; Antsahampano; Antsiranana Rural, -12.323135, 49.294285, 67 m] (Alluaud 1893); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102109 (
Camponotus kelleri
Forel, 1886b: 186. Syntype workers, Madagascar, Toamasina (C. Keller); 1 syntype major worker designated as lectotype, by present designation, AntWeb CASENT0101519 (
Camponotus kelleri var. invalidus
Forel, 1897: 200. Syntype workers, Madagascar, Nosibé [Antsiranana, Nosy be, -13.291667, 48.258335, 146 m] (Voeltzkow); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101517 (
Comoros: Anjouan: Lac Dzialandée, -12.22474, 44.43121, 900 m, disturbed montane rainforest (B.L. Fisher et al.) (
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; two pairs of white spots present on second and third abdominal tergites; pronotum, mesonotum, and propodeum not forming separate convexities, metanotal groove not depressed; propodeum immediately in junction with promesonotum; propodeal dorsum concave.
Minor worker. In full-face view, head sides diverging towards broadly convex posterior margin; eye convex and large (EL/CS: 0.26±0.02; 0.21–0.30), not breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/4 of head (PoOc/CL: 0.25±0.01; 0.22–0.28); frontal carinae widely diverging posteriorly (FR/CS: 0.34±0.01; 0.31–0.37), distance between them larger than their smallest distance to eye; clypeus with anterolateral angle and straight anteromedian margin; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.23±0.07; 1.01–1.36). Promesonotum slightly convex; posterior portion of mesonotum flat immediately anterior to feebly visible metanotal groove; propodeal dorsum immediately posterior to metanotal groove convex, medially strongly concave, then posteriorly flat, joining declivity at a remarkably visible angle; declivity height 3/4 length of dorsum. Petiolar node short and high, with dorsal margin inclined posteriorly and rounding to anterior margin; anterior face shorter than posterior face; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites with a pair of white spots; lateral margin of head lacking erect hairs; two erect hairs present close to posterior margin of head; antennal scape covered only with appressed hairs; pronotum with two and mesonotum with one pairs of erect hairs, respectively; posterodorsal corner of propodeum with a pair of erect hairs. Body color shining reddish black, dark brown to orange-yellow; apical portion of appendages pale in color.
Major worker. Differing from minor worker in the following characters: enlarged subquadrate head (CS: 2.44±0.14; 2.19–2.66; CWb/CL: 1.04±0.03; 1.00–1.08), with broadly concave posterior margin; anteromedian clypeal margin with median excision; antennal scape barely surpassing posterior cephalic margin; strongly built mandible; robust mesosoma, with distinct metanotum; propodeal dorsum slightly to strongly concave towards declivity surface; petiolar node scalelike.
Endemic to the Malagasy region, C. quadrimaculatus is known from Madagascar, the Comoros islands (Anjouan, Grand Comore and Moheli), and Mayotte (Fig.
Camponotus quadrimaculatus A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT0096044 and CASENT0101517 G distribution map.
Camponotus quadrimaculatus is similar to C. rotrae but in the latter the propodeal dorsum is straight.
The minor workers of C. quadrimaculatus display a remarkable range of phenotypic diversity. This assertion of the diversity of morphological characters is made on the basis of specimens with similar promesonotum shape, dorsal outline of the propodeal dorsum, and color of the body. However, the geographic range of differences among the observed variants does not follow a simple pattern. One variant of the yellowish orange body color has a restricted geographic boundary in the north of Madagascar, while others, showing variation in the strength of the concavity on the propodeal dorsum, are widely distributed along the rainforests of the island.
Across the wide geographic range of Camponotus quadrimaculatus, trait variations in the worker castes of C. quadrimaculatus sellaris, Camponotus kelleri and Camponotus kelleri var. invalidus have been observed. However, these variations do not present any pattern that separates them from the former species. Accordingly, C. quadrimaculatus sellaris, Camponotus kelleri and Camponotus kelleri var. invalidus are synonymyzed under Camponotus quadrimaculatus in present study.
Multivariate morphometric analysis through the NC-clustering method was applied to identify the presence of variants detected by the qualitative morphology-based study. However, NC-clustering did not reveal the existence of discrete variants. The samples from each of the variants are spread across the cluster of C. quadrimaculatus. Also, its classification success is only 97.2% because one of these samples was identified by LDA as C. gibber. This might be due to the fact that one of the variants belonging to this species is very similar to C. quadrimaculatus based on the qualitative morphology-based study.
Camponotus hova radamae
Forel, 1891: 31. Lectotype minor worker, by present designation, Madagascar, forêts du versant NE du grand massif (Humblot) AntWeb CASENT0104639 (ZMHB) [examined]. Paralectotypes: with the following data Forêt des bords de l’Ivondrona, près de Tamatave (Dr Conrad Keller) and forêts du versant NE du grand massif (Humblot): 2 minor workers: 1 from Forêt des bords de l’Ivondrona CASENT0101347 (
Madagascar: Antsiranana: 6.5 km SSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.5, 875 m, rainforest (B.L. Fisher) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible closely spaced; lateral cephalic margin posterior to eye level without erect hairs; antennal scape covered with appressed hairs; mesosoma much higher and short, with propodeal dorsum 3 × as long as declivity; petiolar node more or less flattened anteroposteriorly and tapering dorsally.
Minor worker. With head in full-face view, lateral margins anterior to eye level parallel, posteriorly rounding evenly towards short-necked rear margin; eye protruding and large (EL/CL: 0.31±0.01; 0.29–0.34), breaking lateral cephalic margin, location of its posterior margin at posterior 1/3 of head (PoOc/CL: 0.29±0.01; 0.27–0.31); frontal carinae widely opened posteriorly (FR/CS: 0.32±0.01; 0.30–0.34); clypeus with blunt or poorly defined anterolateral angle, anteromedian margin broadly convex; mandible with two apical teeth closely spaced; antennal scape relatively long (SL/CS: 1.52±0.07; 1.42–1.64). Mesosoma with weakly convex promesonotum (MPH/ML: 0.34±0.01; 0.31–0.36), posterior portion flat immediately anterior to metanotal groove; metanotal groove weakly visible, propodeal dorsum almost straight, junction of propodeal dorsum and declivity bluntly angulate, propodeal declivity almost 1/2 length of the dorsum. Petiolar node flattened anteroposteriorly, with dorsal margin rounding to anterior margin; tibia of hind leg rounded axially, without basal twist.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head anterior to level of eyes present but absent behind eye level; antennal scape covered with appressed hairs; posterior margin of head with two erect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 2.68±0.31; 2.29–3.02; CWb/CL: 0.95±0.04; 0.89–0.99) with broadly concave posterior margin; two apical teeth of mandible normally spaced; antennal scape barely surpassing posterior cephalic margin; robust mesosoma, metanotum distinctly visible, propodeal dorsum almost as long as the height of much more vertical declivity, their junction forming a broad angle; more pairs of erect hairs on promesonotum, junction of propodeal dorsum, declivity, and posterodorsal margin of petiolar node.
Endemic to Madagascar, Camponotus radamae normally occurs in eastern humid forests ranging from the littoral region to mountaintops (Fig.
Camponotus radamae A lateral view B head in full-face view C dorsal view of minor worker CASENT0066777 D distribution map.
Camponotus radamae is similar to C. vano in that the two apical teeth of their mandible are closely spaced, but C. vano has a more elongate body in which the mesosoma is very low and long, and its propodeal dorsum is at least four times as long as the declivity surface. In C. radamae the propodeal dorsum is 3 × as long as the declivity.
In Camponotus radamae the species definition is supported by the clustering shown by the dendrogram and confirmed by cumulative LDA at 100% classification success.
Camponotus roeseli
Forel, 1910: 20. Syntype major workers, Madagascar, Montagne d’Ambre (Rolle); 1 syntype major worker designated as lectotype, by present designation, AntWeb CASENT0101602 (
Camponotus maculatus st. legionarium
Santschi, 1911b: 283. Holotype (by monotypy) major worker, Madagascar, Diego Suarez (Légion Étrangère 1903) AntWeb CASENT0101413 (
Madagascar: Antsiranana: [Diego-Suarez]; Museum Paris, Légion Étrangére 1903]; Bevohotra; Sambava, -14, 49.5, 797 m (
In full-face view, lateral cephalic margins converging posteriorly towards eye level, posterior to eye level covered with erect hairs; two apical teeth of mandible closely spaced; head and gaster reddish black and mesosoma reddish orange or integument entirely reddish black.
Minor worker. In full-face view, lateral margins slightly converging progressively towards level of anterior border of eye, strongly converging to a short posterior margin; eye large and broadly convex (EL/CS: 0.27±0.01; 0.25–0.29), breaking lateral cephalic margin, level of its posterior margin located at ca. posterior 1/3 of head (PoOc/CL: 0.28±0.01; 0.26–0.29); frontal carinae medially diverging (FR/CS: 0.24±0.01; 0.23–0.27), posteriorly parallel; clypeus without well-defined anterolateral angle, its anteromedian margin broadly convex, with anterolateral angle and anteromedian margin medially slightly concave; mandible with two apical teeth closely placed; antennal scape relatively long (SL/CS: 1.70±0.08; 1.50–1.81). Pronotum and anterior section of mesonotum weakly convex, posterior portion of mesonotum flat immediately anterior to a barely visible metanotal groove; propodeal dorsum anteriorly convex and posteriorly flat, rounding to declivity; propodeal dorsum ca. 3 × as long as declivity. Petiolar node approximately conical or with short anterior face and posteriorly inclined dorsal margin; femur of hind leg axially rounded, not twisted basally.
First and second gastral tergites without a pair of white spots; head with numerous erect hairs on lateral margin, more than six erect hairs present near its posterior margin; antennal scape covered in suberect hairs inclined at ca. 30°; promesonotum, posterior portion of propodeum, and higher level of declivity covered with more or less long, erect hairs.
Major worker. With characteristics of minor worker except: broader head (CS: 14.07±0.19; 3.81–4.40; CWb/CL: 0.89±0.04; 0.85–0.96), apical 1/3 of antennal scape extending beyond posterior cephalic margin; two apical teeth of mandible normally spaced, usually area ranging from third to basal teeth of mandible divided into two teeth or denticles; mesosoma very robust, promesonotum and metanotum forming even convexity, propodeal dorsum forming separate convexity and rounding to a short declivity surface; petiolar node more flattened anteroposteriorly. Hairs numerous and shorter on body dorsum.
As an endemic and widespread species of Madagascar, C. roeseli generally occurs in the western portion of the island in areas ranging from littoral and rainforest habitats in the north to dry forest areas in the west-central region through spiny forest and thicket in the south (Fig.
Camponotus roeseli A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT0492473 and CASENT0179449 G distribution map.
Camponotus roeseli might be confused with C. daraina because of the presence of erect hairs on the lateral margin of the head posterior to the eye level; however, the latter species can be distinguished by the reddish orange color of the entire body.
The species name C. roeseli and the subspecies name C. maculatus legionarium were created and described by
The taxonomic identity of C. roeseli established by conventional qualitative morphology is supported by multivariate statistical methods. The grouping of samples of this species achieved by the NC-clustering method is confirmed by LDA with an identification success of 100%.
Madagascar: Province Fianarantsoa: PN Isalo, 9.1 km 354° N Ranohira, -22.48167, 45.46167, 725 m, gallery forest, ex rotten log, 27–31 Jan 2003 (Fisher, Griswold et al.) collection code: BLF07331, specimen code: CASENT0485625 (
1 major worker of same data as holotype but specimen coded as: CASENT0485628 (
Madagascar: Fianarantsoa: dry wash, 1 km E of PN Isalo Interpretive Center, -22.62667, 45.35817, 885 m, dry wash (R. Harin’Hala) (
In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; two pairs of white spots present on second and third abdominal tergites; pronotum, mesonotum, and propodeum not forming separate convexities, metanotal groove not depressed; propodeum immediately in junction with promesonotum; propodeal dorsum straight.
Minor worker. In full-face view, lateral margin of head diverging posteriorly towards level of eye, rounding evenly to posterior margin behind eye level; eye protruding and small (EL/CS: 0.24±0.01; 0.22–0.26), not interrupting lateral cephalic border, level of its posterior margin located approximately at posterior 1/4 of head (PoOc/CL: 0.25±0.01; 0.23–0.25); frontal carinae wide (FR/CS: 0.31±0.01; 0.29–0.33), posteriorly diverging, distance between them larger than their smallest distance to eye; clypeus with anterolateral angle and straight anteromedian margin; two apical teeth of mandible normally distant; antennal scape relatively long (SL/CS: 1.19±0.05; 1.08–1.27). Promesonotum weakly convex, mesopropodeum almost flat; metanotal groove barely visible or indistinct; propodeal dorsum nearly straight, its junction to declivity bluntly angulate; propodeal dorsum two and a 1/2 times as long as declivity. Petiolar node short and high, with dorsal margin inclined posteriorly and rounding to anterior margin; anterior and posterior faces at almost the same height; femur of hind leg rounded axially, not twisted near base.
First and second gastral tergites with a pair of white spots; lateral margin of head without erect hairs; two erect hairs present near posterior margin of head; antennal scape covered only with appressed hairs; pronotum with few erect hairs; mesonotum with one pair of erect hairs; posterodorsal corner of propodeum with two pairs of erect hairs. Body color shining brown to dark-brown; apical portion of appendages with lighter color.
Major worker. Differing from minor worker in the following characters: larger head (CS: 2.29±0.17; 2.03–2.42; CWb/CL: 1.03±0.01; 1.02–1.04) with more or less straight posterior margin; apical 1/5 of antennal scape surpassing posterior cephalic margin; in lateral view, mesosoma characterized by even convexity of pronotum and mesonotum, separated from propodeum by distinct metanotum; propodeal dorsum strongly sloping to declivity surface and two and a 1/2 times as long as declivity.
Endemic to Madagascar, C. rotrae is geographically limited to the south-central high plateau of Madagascar. The species occupies montane rainforests of the PN Andringitra, gallery forest and grassland of the PN Isalo, savannah woodland of Itremo, and relict montane rainforest of the Forêt Classée d’Analavelona (Fig.
Camponotus rotrae A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0485625 D distribution map.
See discussion under C. gibber. The qualitative morphological distinction of C. rotrae from other species has been confirmed by multivariate analyses of quantitative morphology. The cluster of C. rotrae samples in the dendrogram of the NC-clustering is supported by LDA with a classification success of 100%.
The species name rotrae is a non-Latin singular noun used in apposition.
Madagascar: Province Antsiranana: RS Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest, ex rotten log, 10–16 Feb 2001, (Fisher, Griswold et al.) collection code: BLF02880, collection code: CASENT0428077 (
3 minor workers of the same data as holotype but with the following specimen codes: CASENT0428078, CASENT0428080, CASENT0428079 (
Madagascar: Antsiranana: Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (
With head in full-face view, lateral cephalic margins converging posteriorly towards eye level; anteromedian margin of clypeus broadly convex; two apical teeth of mandible normally spaced; lateral cephalic margin anterior to eye level without erect hairs.
Minor worker. In full-face view, lateral margins of head converging progressively to level of anterior ocular margin, strongly converging to a short posterior margin behind eye level; eye protruding and large (EL/CS: 0.28±0.01; 0.27–0.29), breaking lateral cephalic border, level of its posterior margin at least at posterior 1/3 of head (PoOc/CL; 0.32±0.04; 0.30–0.42); frontal carinae wide (FR/CS: 0.22±0.0; 0.21–0.23), posteriorly parallel, distance between them larger than smallest distance to eye; clypeus without well-defined anterolateral angle, its anteromedian margin broadly convex or triangular; mandible with two apical teeth closely spaced; antennal scape relatively long (SL/CS: 2.04±0.07; 1.97–2.18). Mesosoma low and long (MPH/ML: 0.33±0.03; 0.28–0.36), with weakly convex promesonotum and approximately flat mesopropodeum; metanotal groove barely visible; propodeal dorsum almost straight or anteriorly slightly convex and posteriorly flat; dorsal margin of propodeum and declivity joined at blunt angle; propodeal declivity 1/3 of length of dorsum. Petiole nodiform, dorsal margin inclined posteriorly, its junction to anterior face bluntly angulate; anterior face of petiolar node 1/2 height of posterior face; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; lateral margin of head without erect hairs; two erect hairs present near posterior cephalic margin; antennal scape without erect hairs but covered with numerous appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum without erect hairs.
Major worker. Differing from minor worker in the following characters: larger head (CS: 4.45; CWb/CL: 0.98); apical 1/4 of antennal scape extending beyond posterior cephalic margin; very robust, short, and high mesosoma, with promesonotum and metanotum forming an even convexity; and propodeal dorsum sloping and joining declivity with blunt angle. Petiolar node much compressed anteroposteriorly, anterior face shorter than posterior.
Camponotus sambiranoensis is only known to occur in the north of Madagascar. It colonizes the dry forests of Binara, RS Ambre, and RS Ankarana, the transitional forests of Ampasindava and Binara, the rainforests of Makirovana and RS Manongarivo, and the montane forest of RS Manongarivo (Fig.
Camponotus sambiranoensis A lateral view B head in full-face view C dorsal view of minor worker CASENT0231845 D distribution map.
Camponotus sambiranoensis shares a broadly convex anteromedian margin of clypeus with C. niavo, and C. cervicalis, but in the two latter species the lateral cephalic margin anterior to the eye level is covered with erect hairs.
The qualitative, morphology-based delimitation of C. sambiranoensis is confirmed by the multivariate morphometric technique. The validity of the species is also confirmed by LDA at 100% classification success.
The species name sambiranoensis is a Latin singular adjective in the nominative case of masculine gender. This species name refers to the region of Sambirano where the species was found.
Camponotus maculatus st. strangulatus
Santschi, 1911a: 129. Lectotype major worker, by present designation, Madagascar, Vitikampy; Morondava, AntWeb CASENT0101097 (
Camponotus hova maculatoides
Emery, 1920b: 6. Lectotype minor worker, by present designation, Madagascar, Nosibe, Antsiranana (Voeltzkow) AntWeb CASENT0101095 (
Note: As junior synonym by Emery 1920: 6. [Emery established Camponotus strangulatus as the junior synonym of Camponotus maculatoides but the former name has priority:
Comoros: Grande Comore: Domani, -11.51778, 43.28, 25 m, coastal scrub (B.L. Fisher et al.) (
With head in full-face view, lateral cephalic margins anterior to eye level parallel, lacking erect hairs; in oblique profile, three pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin; clypeus with distinct anterolateral corner; in profile, junction of propodeal dorsum to declivity rounded; petiole nodelike and not anteroposteriorly compressed.
Minor worker. In full-face view, head sides anterior to eye level parallel, converging strongly towards posterior margin behind eye level; eye convex (EL/CS: 0.28±0.01; 0.26–0.30), breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/3 of head (PoOc/CL: 0.27±0.01; 0.26–0.29); frontal carinae close, distance between them equal to or smaller than their smallest distance to eye (FR/CS: 0.25±0.01; 0.24–0.27); clypeus with anterolateral angle and an almost straight or bluntly angulate anteromedian margin; two apical teeth of mandible normally spaced; antennal scape relatively long (SL/CS: 1.58±0.05; 1.50–1.66). Mesosoma low and long (MPH/ML: 0.33±0.01; 0.31–0.36); promesonotum weakly convex, mesonotum and propodeum nearly flat, separated by weakly visible metanotal groove; propodeal dorsum rounding to declivity; propodeal dorsum roughly 2 × as long as declivity. Petiolar node tapering dorsally, its dorsal margin inclined posteriorly, rounding to anterior margin; anterior face 1/2 height of posterior face; femur of hind leg rounded axially, not twisted near base.
First and second gastral tergites without a pair of white spots; lateral margin of head without erect hairs; a pair of erect hairs present near posterior cephalic margin; with head in profile, three pairs of erect hairs arranged from level of anterior margin of eye to posterior cephalic margin; antennal scape without erect hairs, covered with distantly spaced pubescence; pronotum with two pairs of erect hairs; mesonotum and propodeum without erect hairs.
Major worker. With characteristics of minor worker, except for the following divergent features: enlarged head (CS: 3.22±0.36; 2.73–3.74; CWb/CL: 0.91±0.06; 0.80–0.98), apical 1/4 of antennal scape surpassing posterior cephalic margin, much more robust mesosoma, promesonotum forming a convexity, separated from propodeum by a distinct metanotum; propodeal dorsum convex immediately behind metanotum, length almost equal to height of declivity, their junction at a rounded angle; petiolar node tapering dorsally.
Endemic to the Malagasy region, C. strangulatus occurs in Grand Comore and Moheli of the Comoros islands, in Mayotte, in Aldabra of the Seychelles, and in Madagascar (Fig.
Camponotus strangulatus A lateral view B head in full-face view C dorsal view of minor worker CASENT0132660 D distribution map.
Camponotus strangulatus may be difficult to distinguish from C. tapia and C. atimo but the dorsum of the head in the latter species bears four or more pairs of erect hairs arranged successively from the level of the anterior ocular margin towards the posterior cephalic margin.
As generally known, the genus Camponotus displays polymorphism in the worker castes and this feature is amplified by the phenotypic variation within the castes when the species has specifically a large geographic distribution. This is especially true for C. strangulatus which inhabits most of the terrestrial landscapes, but not the rainforests in the east of Madagascar. Camponotus hova maculatoides, created by
Species delimitation of C. strangulatus on the basis of traditional taxonomic study is congruent with the results of the NC-clustering method. The species was also classified correctly at 100% by confirmatory LDA.
Madagascar: Province Fianarantsoa: Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, grassland, under stone, 22–26 Jan 2003 (Fisher, Griswold et al.) collection code: BLF07214, specimen code: CASENT0493939 (
5 minor workers and 1 major worker with same data as holotype but respectively with specimen codes: CASENT0826114, CASENT0826115, CASENT0493938, CASENT0837568, CASENT0837569, CASENT0493942 (
Madagascar: Antananarivo: Andohony IV Non Protected Area, 22.51 km SW Antsirabe, -20.06718, 46.99274, 1526 m, Uapaca woodland (A. Ravelomanana) (
With head in full-face view, lateral margins of head anterior to eye level parallel, lacking erect hairs; clypeus with distinct anterolateral corner; in oblique profile, four or more pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin; in profile, junction of propodeal dorsum to declivity surface broadly angulate; petiole nodelike and not anteroposteriorly compressed.
Minor worker. In full-face view, head sides anterior to level of eye approximately parallel, strongly converging to form a short area posterior to level of eye; eye protruding and large (EL/CS: 0.29±0.01; 0.28–0.31), breaking lateral cephalic margin, level of its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.24±0.01; 0.23–0.25); frontal carinae close to each other, their distance equal to or smaller than their smallest distance to the eye (FR/CS: 0.25±0.01; 0.23–0.27); clypeus with anterolateral angle, its anteromedian margin broadly triangular or convex; mandible with six teeth, its two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.61±0.08; 1.48–1.71). Promesonotum weakly convex, mesopropodeum almost flat; mesonotum flat immediately anterior to metanotal groove; metanotal groove weakly visible; propodeal dorsum anteriorly slightly convex and posteriorly flat, joining declivity at a blunt angle; propodeal declivity 1/3 length of dorsum. Petiole nodiform, its dorsal margin rounding to anterior face, posterior face higher than anterior; femur of hind leg rounded axially, without twist basally.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head absent; posterior margin of head with two erect hairs; with head in profile, four pairs of erect hairs arranged from level of anterior margin of eye to posterior cephalic margin; antennal scape covered only with appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum with 2–4 erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged head (CS: 3.20±0.21; 2.84–3.38; CWb/CL: 0.99±0.03; 0.94–1.01) with noticeably concave posterior margin; apical 1/3 of antennal scape surpassing posterior cephalic margin; robust mesosoma, propodeal dorsum convex immediately posterior to metanotum, propodeal dorsum < 2 × as long as height of declivity, rounded at junction; petiolar node tapering dorsally.
The distribution of C. tapia is limited to the high central plateau of Madagascar (Fig.
Camponotus tapia A lateral view B head in full-face view C dorsal view of minor worker CASENT0217310 D distribution map.
See discussion under C. atimo.
In the present study, the definition of C. tapia is based on both qualitative morphological analysis and morphometrics, which are congruent. The grouping of the samples of this species obtained from the NC-clustering is confirmed by LDA with an identification success of 100%.
The species name tapia is non-Latin singular noun used in apposition and refers to the Malagasy name for Uapaca in reference to the type of habitat where the ant was most frequently found.
Madagascar: Province Toamasina: RNI Betampona, 34.08 km 332° Toamasina, -17.91977, 49.20039, 525 m, rainforest, Malaise trap, 06 Apr 2008–01 Feb 2009 (B.L. Fisher et al.) collection code: BLF19594, specimen code: CASENT0145284 (
3 minor workers with same data as holotype but specimen coded as: CASENT0138778, CASENT0138799, CASENT0138800 (
Madagascar: Antsiranana: Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 650–800 m, rainforest (B.L. Fisher et al.) (
With head in full-face view, lateral cephalic margins converging posteriorly towards eye level and covered with erect hairs; erect hairs lacking posterior to eye level; anteromedian margin of clypeus broadly convex; two apical teeth of mandible normally spaced.
Minor worker. In full-face view, head sides anterior to level of eye parallel; lateral cephalic margins posterior to level of eye converging progressively to posterior margin; eye large (EL/CS: 0.28±0.01; 0.27–0.29), protruding, and breaking lateral cephalic border, level of posterior margin located at posterior 1/3 of head (PoOc/CL: 0.31±0.02; 0.30–0.33); frontal carinae slightly diverging posteriorly (FR/CS: 0.25±0.01; 0.24–0.26), larger than smallest distance to eye; clypeus with rounded anterolateral corner and broadly angulate anteromedian margin; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.95±0.00; 1.95–1.96). Mesosoma low and long (MPH/ML: 0.30); dorsum of mesosoma more or less flat, promesonotum weakly convex, posterior portion of mesonotum flat immediately anterior to barely visible metanotal groove; propodeal dorsum anteriorly slightly convex and posteriorly flat, its junction with declivity bluntly angulate; propodeal dorsum 3 × as long as declivity. Petiole nodiform, with dorsal margin inclined posteriorly and forming a blunt angle to anterior margin; anterior face 1/2 height of posterior face, which inclines anteriorly to the dorsal face; femur of hind leg rounded axially, not twisted basally.
First and second gastral tergites without a pair of white spots; lateral margin of head anterior to level of eyes covered with erect hairs, which are lacking posterior to level of eyes; two erect hairs present near posterior cephalic margin; antennal scape covered with suberect hairs inclined at ca. 30° as well as appressed hairs; pronotum with a pair of erect hairs; posterodorsal angle of propodeum with a pair of erect hairs.
Major worker. Unknown.
The distribution of C. tendryi is limited to Madagascar, in rainforest habitats of the RNI Betampona and RS Ambatovaky in the east, and the transitional humid forest of Binara in the northeast (Fig.
Camponotus tendryi A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0145284 D distribution map.
See discussion under C. dufouri.
Camponotus tendryi were not included in the morphometric analysis due to the lack of sufficient minor worker samples.
The species name tendryi is a Latin noun in the genitive case derived from the first name of a male person “Tendry”.
Madagascar: Province Toamasina: PN Zahamena, Onibe River, -17.75908, 48.85468, 780 m, rainforest, beating low vegetation, 22 Feb 2009 (B.L. Fisher, Malagasy Arthropod Team) collection code: BLF22341, specimen code: CASENT0217316 (
2 minor workers and 1 major worker with same data as holotype but respectively specimen coded as: CASENT0077651, CASENT0840785, CASENT0152085 (
Madagascar: Toamasina: PN Zahamena, Onibe River, -17.75908, 48.85468, 780 m, rainforest (B.L. Fisher, Malagasy Arthropod Team) (
Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandibles closely spaced; lateral cephalic margin posterior to eye level without erect hairs; antennal scape covered with appressed hairs; mesosoma very low and long; propodeal dorsum at least four times as long as declivity; petiole nodelike.
Minor worker. With head in full-face view, lateral margins anterior to eye level parallel, posteriorly rounding evenly towards a rear margin that is somewhat necked; eye protruding and large (EL/CL: 0.29), breaking lateral cephalic margin, location of its posterior margin at posterior 1/3 of head (PoOc/CL: 0.34); frontal carinae widely opened (FR/CS: 0.29±0.00; 0.29–0.30), posteriorly parallel; clypeus with blunt or poorly defined anterolateral angle, anteromedian margin broadly convex; two apical teeth of mandible closely spaced; antennal scape relatively long (SL/CS: 1.68±0.03; 1.66–1.70). Mesosoma low and long (MPH/ML: 0.29±0.02; 0.28–0.31), promesonotum slightly convex, mesonotum and propodeum more or less flat; metanotal groove feebly visible, propodeal dorsum rounding to declivity, propodeal dorsum four times as long as declivity. Petiole nodelike, with dorsal margin inclined posteriorly and rounding to anterior margin; tibia of hind leg rounded axially, without basal twist.
First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head present anterior to eye level and absent behind eye level; posterior margin of head with two erect hairs; antennal scape covered with appressed hairs; junction of propodeal dorsum and declivity with a pair of erect hairs.
Major worker. Differing from minor worker in the following characters: enlarged, elongate, and rectangular head (CS: 1.67; CWb/CL: 0.64) with slightly concave posterior margin; two apical teeth of mandible normally spaced; distal 1/3 of antennal scape surpassing posterior cephalic margin; robust, short, and high mesosoma, metanotum distinctly visible, propodeal dorsum almost 3 × as long as more vertical declivity, their junction forming a blunt angle; with more pairs of erect hairs. Head dark brown, body color pale brown, propodeum much darker.
A species endemic to Madagascar, C. vano is geographically restricted to the PN Zahamena and the Corridor Forestier Analamay-Mantadia in humid forests in the east of the island (Fig.
Camponotus vano A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217316 D distribution map.
See discussion under C. radamae.
Camponotus vano was not included in the NC-clustering analysis due to an inadequate number of minor workers. However, its qualitative morphological traits are sufficient to supports its status as a real species.
The species name vano is a non-Latin singular noun used in apposition and refers to the Malagasy name for “heron”. It refers to the elongate body form of the species.
We would like to express our gratitude to B. Landry from
Table S1
Data type: Morphological measurements
Explanation note: Table S1. Basic measurements of individual specimens arranged by species code, collection code, and specimen code (unique identification number). See text for abbreviations.