Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology

Jean Claude Rakotonirina; Brian L. Fisher

doi:10.3897/zookeys.1098.73223

Research Article

Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology

Jean Claude Rakotonirina^‡§, Brian L. Fisher^|

‡ Madagascar Biodiversity Center, Antananarivo, Madagascar

§ Université d’Antananarivo, Antananarivo, Madagascar

| Entomology, California Academy of Sciences, San Francisco, United States of America

Corresponding author: Jean Claude Rakotonirina ( jcrakoto25@yahoo.com )

Academic editor: Marek Borowiec

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

ZooBank: urn:lsid:zoobank.org:pub:B4F4033F-296E-43CC-BE54-B9413BC19268

Abstract

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.

Keywords

Madagascar, morphometry, species delimitation, subgenus Myrmosaga, taxonomy

Introduction

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 (Rakotonirina et al. 2016, 2017; Rasoamanana et al. 2017; Rakotonirina and Fisher 2018). We continue to apply these techniques to revise the subgenus Myrmosaga (Forel 1912) as part of an ongoing project to revise the entire genus in the Malagasy region.

Myrmosaga was created by Forel (1912) as a subgenus of Camponotus, but later it was synonymized under Mayria by Emery (1925) based on a few morphological characters that are shared by both subgenera. However, our observation of the samples collected from the recent extensive inventory of ant fauna in the Malagasy region revealed two separate groups. One group includes the species of Myrmosaga and another contains species that remain in the group Mayria. In addition, Camponotus imitator, the type species of the subgenus Myrmopytia (Emery, 1920a) is grouped with the species of Myrmosaga. This grouping is also supported by the molecular analysis using UCE (Ultra-Conserved Element) phylogenomic data (Unpublished data). Species of Myrmosaga are remarkably different from those of Mayria and therefore the subgenus needs to be revived now from synonymy. Myrmopytia is also synonymized under Myrmosaga in the present study. The other three species (C. jodina, C. karaha, and C. longicolis) recently described under Myrmopytia will be moved to a different subgenus (Rasoamanana in prep.).

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 1).

Table 1.

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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.

Materials and methods

Abbreviation of depositories

CAS California Academy of Sciences, San Francisco, CA, USA.

MHNG Musée d’Histoire Naturelle, Geneva, Switzerland.

MNHN Musée National d’Histoire Naturelle, Paris, France.

MSNG Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy.

NHMB Naturhistorisches Museum, Basel, Switzerland.

NHMUK NNatural History Museum, London, UK.

PBZT Parc Botanique et Zoologique de Tsimbazaza, Antananarivo, Madagascar.

PSWC P.S. Ward Collection, University of California at Davis, CA, USA.

ZMHB Museum für Naturkunde der Humboldt Universität, Berlin, Germany.

Materials

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 1). Collection event codes with prefix AND, ANTC, ARA, BLF, MG, or PSW indicate distinct collecting events, and were used as grouping factors in the NC-clustering method. One grouping factor may contain up to three individual specimens for measurements.

Methods

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 (Hijmans et al. 2011). The major ecoregions of Madagascar were superimposed on the distribution of each species. Specimens with inadequate geographic coordinates were excluded from these maps. Information about the biology of each species was based on data obtained from specimen sampling (the nest series and isolated worker specimens) in the field.

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).

Measurements

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 Rakotonirina et al. 2016: fig. 1).

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.

Multivariate statistical analyses

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 1), and (2) the ratios (indices) of measurements involving the comparison of one measured trait (variable) over another (CS, CL, or ML) to show the body proportions or shape of the specimen (Table 2). In Table 2, the ratios of measurements for minor and major workers of each species are presented as arithmetic means with standard deviation in the upper line and as ranges in square brackets in the lower line if more than two individual specimens are measured.

Table 2.

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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
aina	Minor (n= 2)	[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
	Minor (n= 31)	[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
	Major (n= 5)	[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
	Minor (n= 19)	[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
	Major (n= 4)	[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
	Minor (n= 31)	[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
	Major (n= 8)	[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
	Minor (n= 16)	[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
	Major (n= 4)	[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
	Minor (n= 11)	[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
	Major (n= 3)	[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
bemaheva	Minor (n= 23)	[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
	Minor (n= 49)	[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
	Major (n= 13)	[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
	Minor (n= 12)	[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
	Major (n= 6)	[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
	Minor (n= 9)	[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
	Major (n= 7)	[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
	Minor (n= 8)	[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
	Major (n= 5)	[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
	Minor (n= 19)	[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
	Major (n= 5)	[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
	Minor (n= 19)	[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
	Major (n= 5)	[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
	Minor (n= 37)	[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
	Major (n= 9)	[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
	Minor (n= 18)	[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
	Major (n= 6)	[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
	Minor (n= 17)	[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
	Major (n= 5)	[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
harenarum	Minor (n= 5)	[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
	Minor (n= 27)	[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
	Major (n= 10)	[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
	Minor (n= 55)	[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
	Major (n= 17)	[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
imitator	Minor (n= 44)	[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
	Minor (n= 12)	[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
	Major (n= 7)	[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
joany	Minor (n= 2)	[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
karsti	Minor (n= 2)	[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
	Minor (n= 17)	[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
	Major (n= 7)	[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
	Minor (n= 44)	[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
	Major (n= 12)	[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
lokobe	Minor (n= 8)	[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
mahafaly	Minor (n= 19)	[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
	Minor (n= 25)	[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
	Major (n= 8)	[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
	Minor (n= 8)	[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
	Major (n= 7)	[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
	Minor (n= 72)	[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
	Major (n= 8)	[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
	Minor (n= 24)	[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
	Major (n= 7)	[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
	Minor (n= 29)	[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
	Major (n= 10)	[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
	Minor (n= 18)	[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
	Major (n= 4)	[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
	Minor (n= 8)	[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
	Minor (n= 18)	[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
	Major (n= 6)	[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
	Minor (n= 17)	[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
	Major (n= 5)	[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
tendryi	Minor (n= 2)	[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
	Minor (n= 2)	[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 Rakotonirina et al. 2016, 2017), we applied morphometric data obtained from minor workers for multivariate statistical analysis.

Species hypotheses were generated using the Nest Centroid clustering (NC-clustering) technique. The procedure followed Rakotonirina et al. (2016).

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.

Results and discussion

Multivariate statistical analysis of morphometric data

The NC-clustering dendrogram revealed 33 clusters, which are interpreted as 33 species (Fig. 1). The 33 species hypothesis proposed by the exploratory analysis was supported by the cumulative LDA, with 98.2% identification success (Table 3). Few species have conflicting identification features. These misidentifications could be attributed to similar morphology. Morphologically similar species frequently exhibit overlapping ranges of quantitative measurements and share qualitative morphological characters (Table 3).

Table 3.

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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
Observed 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

Figure 1.

Dendrogram of NC-Clustering of the subgenus Myrmosaga. Label on the tip of the branch indicates the species name followed by the specimen code.

Taxonomic synopsis of the subgenus Myrmosaga

Myrmosaga Forel 1912: 92 [as subgenus of Camponotus]. Type species: Camponotus kelleri Forel, 1886b: 186, by subsequent designation of Wheeler 1913: 81, a junior synonym of Camponotus quadrimaculatus Forel, 1886a: cii. [As junior synonym of Mayria Emery, 1925: 121]. Stat. rev.

Myrmopytia Emery, 1920a: 243 [as subgenus of Camponotus]. Type species: Camponotus imitator Forel, 1891: 209, by monotypy, by original designation. Syn. nov.

Morphological diagnosis of the worker caste of Malagasy Myrmosaga

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.

Minor worker

Head elongate in full-face view, sides either converging or diverging posteriorly towards eye level to approximately parallel to each other and start rounding to posterior margin at ca. posterior 1/3 to posterior 1/5; posterior margin sometimes indistinct from lateral margins.
Mandible triangular, apical margin armed with six teeth. In some species, two apical teeth closer to each other than the other teeth.
Palp formula: 6,4.
Clypeus with median carina, its anteromedian margin mostly broadly convex, sometimes straight (C. dufouri, C. lubbocki, C. gibber, C. rotrae, C. immaculatus, etc.), projecting into triangular lobe (C. sambiranoensis, C. liandia); anterior margin rarely excised medially (C. lokobe).
Antenna with 12 antennomeres, elongate flagellomeres; antennal scape long, generally its distal 1/2 surpassing posterior cephalic margin, either covered or without erect hairs; apical antennomere slightly longer than other flagellomeres;
Frontal lobe narrow and partially covering the antennal insertion; frontal carina S-shaped, strongly divergent posteriorly.
Compound eye large, most often protruding, its posterior margin usually located at ca. posterior 1/3 to posterior 1/4 of the head.
Mesosoma in lateral view, ranging from long and low (C. dufouri, C. lokobe, C. atimo, C. vano, etc.) to short and high (C. cemeryi); or with pronotum and mesonotum weakly convex, mesonotum and propodeum almost straight (C. hovahovoides).
Promesonotal suture visible.
Pronotum with rounded junction between its dorsum and lateral face; humeral angle rounded.
Mesopleuron and propodeal surface together clearly longer than lateral portion of pronotum in lateral view.
Metapleural gland absent.
Metanotal groove inconspicuous to clearly visible.
Procoxa of normal size, maximum width as large as, or smaller than, width of mesopleuron.
Middle and hind tibiae with single pectinate spur.
Tibia of hind leg either axially rounded or rarely twisted basally.
Petiolar node laterally narrow and low.
Gaster generally elongate and narrow, anteriorly low and short.
Sculpture varying from smooth and shiny through finely and densely imbricate to generally matte.

Major worker

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.

Comments

Originally Forel (1912) created Myrmosaga as a separate subgenus of Camponotus. Its synonymy under Mayria by Emery (1925) was based on similarity in the form of the clypeus, number of mandibular teeth, and shape of the mesosoma, and the petiole. It also was based on insufficient samples and comparative studies of different species belonging to these two subgenera. In the present study, we revealed many characters (in the Morphological diagnosis above) to distinguish Myrmosaga from Mayria. Members of Mayria are differentiated by the combination of the following characters: the clypeus lacks median carina, its anterolateral corner is rounded; the anterior margin of the pronotum is strongly convex in lateral view, forming a rounded flange and extended laterally to form an obtuse humeral angle; in dorsal view, the pronotal disc is rectangular with distinct lateral margins; the propodeum dorsum is never concave. Thus, Myrmosaga is revived from this synonymy here.

Species of the Malagasy Camponotus that have been previously placed in the subgenera Dinomyrmex and Myrmoturba by Forel (1914), in the subgenus Tanaemyrmex by Emery (1925), and two species (C. liandia and C. lubbocki) placed in the subgenus Mayria (Santschi 1914; Rakotonirina and Fisher 2018) are now moved into the subgenus Myrmosaga. This is because our detailed examination of the samples of Camponotus from the recent extensive survey of ants in Madagascar revealed that morphological differences between these taxa are due to their ability to adapt to diverse habitats. Similarities among them are the result of convergence by using similar techniques in exploiting similar habitats.

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 (Rasoamanana et al. 2017). They are specifically characterized by the elongate mesonotum, which is constricted at midlength, and by the dorsally protruding rounded propodeum. However, C. imitator has a medially carinate clypeus with a broad rectangular projection, and a dorsally tapering petiolar node that are considered as some of the strong characteristics of the subgenus Myrmosaga. The other three species have triangular anterior clypeal margin and a lower anterior face and conical petiole. The specific shape of the mesosoma is just a result of the adaptive radiation among these three morphologically similar species across the terrestrial landscapes in Madagascar. Accordingly, C. imitator is presently moved to Myrmosaga and thus Myrmopytia is synonymized under the same subgenus. The three other species recently described under Myrmopytia are moved to a different subgenus (Rasoamanana in prep.).

Synoptic list of species of the Malagasy Myrmosaga

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 4).

Table 4.

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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

Identification key to minor worker caste of the Malagasy Camponotus subgenus Myrmosaga

1	With head in full-face view, eyes not breaking lateral cephalic margins (Fig. 2A)	2
–	With head in full-face view, eyes breaking lateral cephalic margins (Fig. 2B), if not then lateral cephalic margin diverging posteriorly (Fig. 2C)	5
2	Dorsal margin of propodeum entirely straight (Fig. 3A)	3
–	Dorsal margin of propodeum with blunt angle at ca. posterior 1/2 (Fig. 3B)	4
3	Propodeal dorsum 3 × as long as the declivity; petiolar node with dorsal margin as long as posterior margin (Fig. 4A)	*aina*
–	Propodeal dorsum 2 × as long as the declivity; petiolar node with dorsal margin shorter than posterior margin (Fig. 4B)	*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. 5A)	*karsti*
–	With mesosoma in lateral view, mesonotum and anterior 1/2 of propodeum forming separate convexities; petiolar node ca. as high as long (Fig. 5B)	*harenarum*
5	With head in full-face view, lateral cephalic margins converging posteriorly towards eye level (Fig. 6A)	6
–	With head in full-face view, lateral cephalic margin either approximately parallel or diverging posteriorly towards eye level (Fig. 6B, C)	14
6	Two apical teeth of mandible normally spaced (Fig. 7A); clypeus with anterolateral corner	7
–	Two apical teeth of mandible closely spaced (Fig. 7B); clypeus without anterolateral corner, lateral and anteromedian margin continuously forming broad convexity	12
7	With head in full-face view, anteromedian margin of clypeus broadly convex (Fig. 8A)	8
–	With head in full-face view, anteromedian margin of clypeus either generally straight (Fig. 8B) or medially excised (Fig. 8C)	10
8	With head in full-face view, lateral cephalic margin anterior to eye level without erect hairs (Fig. 9A)	*sambiranoensis*
–	With head in full-face view, lateral cephalic margin anterior to eye level covered with erect hairs (Fig. 9B)	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. 10A)	*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. 10B)	*cervicalis*
10	Anteromedian margin of clypeus noticeably excised medially (Fig. 11A)	*lokobe*
–	Anteromedian margin of clypeus straight (Fig. 11B)	11
11	Lateral cephalic margin posterior to eye level covered with erect hairs (Fig. 12A); larger size (CS: 1.97±0.19; 1.66–2.26; ML: 4.23±0.39; 3.56–4.81)	*dufouri*
–	Lateral cephalic margin posterior to eye level without erect hairs (Fig. 12B); smaller size (CS: 1.57±0.01; 1.56–1.58; ML: 3.26±0.01; 3.21–3.31)	*tendryi*
12	Lateral cephalic margin posterior to eye level without erect hairs (Fig. 13A)	*bemaheva*
–	Lateral cephalic margin posterior to eye level covered with erect hairs (Fig. 13B)	13
13	Integument of entire body yellowish orange to reddish orange (Fig. 14A)	*daraina*
–	Head and gaster reddish black and mesosoma reddish orange or integument entirely reddish black (Fig. 14B)	*roeseli*
14	In full-face view, lateral margin of head anterior to eye level approximately parallel and covered with erect hairs (Fig. 15A)	15
–	In full-face view, lateral margin of head anterior to eye level diverging posteriorly (Fig. 15C), if parallel then lacking erect hairs (Fig. 15B)	23
15	With mesosoma in lateral view, mesonotum elongate and constricted at midlength, propodeum broadly convex (Fig. 16A)	*imitator*
–	With mesosoma in lateral view, mesonotum short and lacking constriction at midlength; promesonotum an even convexity; propodeal dorsum almost straight (Fig. 16B)	12
16	Two apical teeth of mandible closely spaced (Fig. 17A)	17
–	Two apical teeth of mandible normally spaced from each other (Fig. 17B)	19
17	Antennal scape covered with erect hairs; lateral cephalic margin posterior to eye level with erect hairs (Fig. 18A); larger in size	*hova*
–	Antennal scape covered with appressed hairs (Fig. 18B); lateral cephalic margin posterior to eye level without erect hairs; smaller in size	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. 19A)	*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. 19B)	*vano*
19	Antennal scape covered with appressed hairs (Fig. 20A)	*mahafaly*
–	Antennal scape covered with erect to suberect hairs (Fig. 20B)	20
20	Mesosoma generally short and high, its dorsal outline strongly convex, propodeal dorsum < 2 × height of declivity surface (Fig. 21A)	*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. 21B)	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. 22A)	*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. 22B)	22
22	Antennal scape covered with suberect hairs inclined at ca. 30° (Fig. 23A); larger species (CS: 1.59±0.10; 1.36–1.75; ML: 3.10±0.16; 2.76–3.45)	*mixtellus*
–	Antennal scape covered with suberect hairs inclined at ca. 45° (Fig. 23B); smaller species (CS: 1.15±0.06; 1.00–1.27; ML: 2.11±0.10; 1.87–2.28)	*boivini*
23	With head in full-face view, lateral margins of head anterior to eye level parallel to each other (Fig. 24A)	24
–	With head in full-face view, lateral margins of head anterior to eye level diverging posteriorly (Fig. 24B)	31
24	In full-face view, clypeus with distinctly visible anterolateral corner (Fig. 25A)	25
–	In full-face view, clypeus without visible anterolateral corner, lateral and anteromedian clypeal margin continuously forming broad convexity (Fig. 25B)	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. 26A)	26
–	Pale yellow to orange body color; in lateral view, petiole nodelike and not compressed anteroposteriorly (Fig. 26B)	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. 27A)	*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. 27B)	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. 28A)	*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. 28B)	*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. 30A)	*atimo*
–	With mesosoma in profile, junction of propodeal dorsum and declivity surface broadly angulate (Fig. 30B)	*tapia*
30	In full-face view, posterior portion of head extending into broadly short neck (Fig. 31A); in profile, entire propodeal dorsum more or less straight and separated from declivity surface by broad angle	*gouldi*
–	In full-face view, posterior portion of head normally rounded, not extending into a short neck (Fig. 31B); in profile, propodeal dorsum immediately posterior to metanotal groove convex, then concave medially and rounding to declivity surface	*aro*
31	With head in full-face view, lateral margin of head anterior to eye level with erect hairs (Fig. 32A); head and gaster black, mesosoma reddish orange to brown	32
–	With head in full-face view, lateral margin of head anterior to eye level without erect hairs (Fig. 32B); body color uniquely black, brown to yellowish orange	33
32	With head in full-face view, anterior clypeal margin broadly triangular (Fig. 33A); mesosoma in profile short and high; gastral tergites covered with abundant pubescence	*aurosus*
–	With head in full-face view, anterior clypeal margin truncate (Fig. 33B); mesosoma in profile low and long; gastral tergites without abundant pubescence	*hagensii*
33	With head in full-face view, anterior clypeal margin broadly triangular (Fig. 34A)	*liandia*
–	With head in full-face view, anterior clypeal margin truncate (Fig. 34B)	34
34	No white spot on dorsum of second and third abdominal tergites (Fig. 35A)	35
–	One pair of white spots on third abdominal tergites (Fig. 35B)or two pairs of white spots present on second and third abdominal tergites (Fig. 35C)	37
35	With mesosoma in lateral view, propodeal dorsum broadly concave (Fig. 36A)	*immaculatus*
–	With mesosoma in lateral view, propodeal dorsum approximately straight (Fig. 36B)	36
36	Eye small (EL/CL: 0.19±0.01; 0.16–0.21) (Fig. 37A); posterior margin of head approximately straight	*kelimaso*
–	Eye large (EL/CL: 0.24±0.01; 0.23–0.27) (Fig. 37B); posterior margin of head broadly convex	*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. 38A)	*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. 38B)	38
38	Propodeal dorsum straight (Fig. 39A)	*rotrae*
–	Propodeal dorsum concave (Fig. 39B)	*quadrimaculatus*

Figure 2.

Head in full-face view A C. joany (CASENT0408908) B C. radamae (CASENT0066777) C C. immaculatus (CASENT0179441).

Figure 3.

Mesosoma in lateral view A C. aina (CASENT0217291) B C. harenarum (CASENT0499207).

Figure 4.

Mesosoma and petiolar node in lateral view A C. aina (CASENT0217291) B C. joany (CASENT0408908).

Figure 5.

Mesosoma and petiolar node in lateral view A C. karsti (CASENT0217292) B C. harenarum (CASENT0499207).

Figure 6.

Head in full-face view A C. sambiranoensis (CASENT0231845) B C. radamae (CASENT0066777) C C. immaculatus (CASENT0179441).

Figure 7.

Mandibles in frontal view A C. niavo (CASENT0300086) B C. bemaheva (CASENT0153778).

Figure 8.

Head in full-face view A C. cervicalis (CASENT0217303) B C. dufouri (CASENT0487727) C C. lokobe (CASENT0436584).

Figure 9.

Head in full-face view A C. sambiranoensis (CASENT0231845) B C. cervicalis (CASENT0217303).

Figure 10.

Head in full-face view A C. niavo (CASENT0134004) B C. cervicalis (CASENT0217303).

Figure 11.

Head in full-face view A C. lokobe (CASENT0436584) B C. tendryi (CASENT0145284).

Figure 12.

Head in full-face view A C. dufouri (CASENT0274127) B C. tendryi (CASENT0145284).

Figure 13.

Head in full-face view A C. bemaheva (CASENT0154158) B C. roeseli (CASENT0492473).

Figure 14.

Body in lateral view A C. daraina (CASENT0077433) B C. roeseli (CASENT0492473).

Figure 15.

Head in full-face view A C. mahafaly (CASENT0115287) B C. tapia (CASENT0493939) C C. quadrimaculatus (CASENT0096044).

Figure 16.

Mesosoma in lateral view A C. imitator (CASENT0452849) B C. mahafaly (CASENT0115287).

Figure 17.

Mandibles in frontal view A C. radamae (CASENT0228250) B C. boivini (CASENT0477264).

Figure 18.

Head in full-face view A C. hova (CASENT0055710) B C. radamae (CASENT0217307).

Figure 19.

Mesosoma and petiolar node in lateral view A C. radamae (CASENT066777) B C. vano (CASENT0217316).

Figure 20.

Head in full-face view A C. mahafaly (CASENT0115287) B C. hovahovoides (CASENT0487242).

Figure 21.

Mesosoma and petiolar node in lateral view A C. cemeryi (CASENT0217306) B C. hovahovoides (CASENT0496908).

Figure 22.

Mesosoma in lateral view A C. hovahovoides (CASENT0496908) B C. mixtellus (CASENT0132606).

Figure 23.

Head in full-face view A C. mixtellus (CASENT0076675) B C. boivini (CASENT0168345).

Figure 24.

Head in full-face view A C. atimo (CASENT0454042) B C. quadrimaculatus (CASENT0096044).

Figure 25.

Head in full-face view A C. tapia (CASENT0217310) B C. aro (CASENT0489917).

Figure 26.

Posterior portion of mesosoma, petiolar node, and gaster in lateral view A C. bozaka (CASENT0217309) B C. atimo (CASENT0454042).

Figure 27.

Head in full-face view A C. becki (CASENT0071380) B C. asara (CASENT0493252).

Figure 28.

Mesosoma and petiolar node in lateral view A C. asara (CASENT0493252) B C. bozaka (CASENT0217309).

Figure 29.

Head in lateral view A C. strangulatus (CASENT0136581) B C. tapia (CASENT0493939).

Figure 30.

Mesosoma in lateral view A C. atimo (CASENT0454042) B C. tapia (CASENT0217310).

Figure 31.

Head in full-face view A C. gouldi (CASENT0121617) B C. aro (CASENT0489917).

Figure 32.

Head in full-face view A C. aurosus (CASENT0064815) B C. gibber (CASENT0188619).

Figure 33.

Head in full-face view A C. aurosus (CASENT0064815) B C. hagensii (CASENT0191568).

Figure 34.

Head in full-face view A C. liandia (CASENT0491145) B C. kelimaso (CASENT0487718).

Figure 35.

Second and third abdominal tergites in dorsal view A C. kelimaso (CASENT0487718) B C. quadrimaculatus (CASENT0096044) C C. gibber (CASENT0491497).

Figure 36.

Head, mesosoma and petiolar node in lateral view A C. immaculatus (CASENT0179441) B C. lubbocki (CASENT0486998).

Figure 37.

Head in full-face view A C. kelimaso (CASENT0487718) B C. lubbocki (CASENT0486998).

Figure 38.

Mesosoma and petiolar node in lateral view A C. gibber (CASENT0188619) B C. quadrimaculatus (CASENT0096044).

Figure 39.

Mesosoma in lateral view A C. rotrae (CASENT0485625) B C. quadrimaculatus (CASENT0121629).

Species accounts

Camponotus aina sp. nov.

http://zoobank.org/B622AD1C-1005-44D3-97F5-05E22C8D291B

Figs 3A, 4A, 40

Holotype worker

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 (CAS).

Paratype

1 minor worker with same data as holotype but specimen coded as: CASENT0217290 (CAS).

Additional material examined

Madagascar: Antsiranana: RS Ankarana, 7 km SSE Matsaborimanga, -12.91667, 49.1, 150 m, tropical dry forest (P.S. Ward) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 40D). Its nest site is unknown, but workers have been found foraging on lower vegetation.

Figure 40.

Camponotus aina A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217291 D distribution map.

Discussion

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.

Etymology

The species name aina is a non-Latin singular noun and is being used as a noun in apposition.

Camponotus aro sp. nov.

http://zoobank.org/3C81BF3C-27C7-476A-889D-F2DD66EE7E55

Figs 25B, 31B, 41

Holotype worker

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 (CAS).

Paratypes

2 minor workers and 1 major worker with same data as holotype but with specimen codes: CASENT0837586, CASENT0837587, CASENT0489918 (NHMUK, MHNG, CAS).

Additional material examined

Madagascar: Mahajanga: PN Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

Geographically restricted to the PN Namoroka in the western dry forest of Madagascar (Fig. 41D), C. aro has been found nesting under stones while workers forage on the ground or through leaf litter and on lower vegetation.

Figure 41.

Camponotus aro A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0489917 D distribution map.

Discussion

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.

Etymology

The species name aro is a non-Latin singular noun used in apposition.

Camponotus asara sp. nov.

http://zoobank.org/5C3637B8-D025-44D1-B91D-E49FA3D3CEBC

Figs 27B, 28A, 42

Holotype worker

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 (CAS).

Paratypes

3 workers, same data as holotype but with the following specimen codes: CASENT0493251, CASENT0837576, CASENT0837577 (NHMUK, CAS).

Additional material examined

Madagascar: Toliara: 15 km E Sakaraha, -22.9, 44.68333, 760 m, tropical dry forest (P.S. Ward) (CAS); PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CAS); RS Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala, -18.26, 45.41833, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); RS Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, -18.26667, 45.40667, 1050 m, montane rainforest (Fisher, Griswold et al.) (CAS); Southern Isoky-Vohimena Forest, -22.68333, 44.83333, 730 m (Sylvain) (CAS); Vohibasia Forest, 59 km NE Sakaraha, -22.46667, 44.85, 780 m (Sylvain) (CAS); PN Zombitse, 19.8 km 84° E Sakaraha,-22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CAS); near ANGAP office, PN Zombitse, -22.8865, 44.69217, 840 m, deciduous spiny forest (R. Harin’Hala) (CAS); near road, PN Zombitse, -22.8405, 44.73117, 825 m, spiny deciduous forest (R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 42D). Colony nests can be in rotten logs and in rot pockets above the ground. Workers have been found foraging in leaf litter and on the ground.

Figure 42.

Camponotus asara A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0493252 D distribution map.

Discussion

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%.

Etymology

The species name asara is a non-Latin singular noun used in apposition.

Camponotus atimo sp. nov.

http://zoobank.org/29081C2B-DA66-41BE-9E62-EF67D84D226B

Figs 24A, 26B, 30A, 43

Holotype worker

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 (CAS).

Paratype

1 major worker same data as holotype but specimen coded as: CASENT0454043 (major) (CAS).

Additional material examined

Madagascar: Antsiranana: Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest, (J.-J. Rafanomezantsoa et al.) (CAS); RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Sahamalaza Peninsula, Forêt d’Anabohazo, 21.6 km 247° WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest (Fisher, Griswold et al.) (CAS). Fianarantsoa: Forêt d’Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, Uapaca woodland (Fisher, Griswold et al.) (CAS). Mahajanga: Melaky Region, District of Maintirano, Ampasy 50 km E of Maintirano, -18.004, 44.452, 85 m, dry forest (Mike, Rin’ha) (CAS). Toliara: 3.4 km 190° S Marovato, -25.55972, 45.2825, 160 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 3.5 km 236° SW Marovato, -25.55389, 45.25583, 230 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 4.4 km 148° SSE Lavanono, -25.45056, 44.97417, 60 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 7.0 km 156° SSE Lavanono, -25.47111, 44.9885, 50 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Anosy Region, District of Amboasary, PN Andohahela, Parcelle III, Ihazofotsy, 32 km NE Amboasary, -24.83083, 46.53617, 58 m, dry forest, spiny forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, District of Fort-Dauphin, PN Andohahela, Parcelle II, Tsimela, 42 km W of Fort-Dauphin, -24.93683, 46.62667, 176 m, transition forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82466, 46.60111, 100 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo Andrefana Region, District of Betioky ; RS Beza Mahafaly Parcelle Belle vue 07 km W of Research Station, -23.68983, 44.5755, 177 m, spiny forest, (Rin’ha) (CAS); Atsimo-Andrefana Region, -23.55275, 43.74471, 45 m, coastal scrub on limestone (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo-Andrefana Region, -23.53922, 43.77935, 20 m, riparian scrub (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo-Andrefana Region, Antsokay Arboretum, -23.41491, 43.75499, 13 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Forêt de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Mahavelo, Isantoria River, -24.75833, 46.15717, 110 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Forêt de Tsinjoriaky, 6.2 km 84° E Tsifota, -22.80222, 43.42067, 70 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Mahafaly Plateau, 6.2 km 74° ENE Itampolo, -24.65361, 43.99667, 80 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Andohahela, Forêt d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, -24.93, 46.6455, 300 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, -23.99222, 43.88067, 90 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Mitoho Cave, 6.4 km 77° ENE Efoetse, 17.4 km 170° S Beheloka, -24.04722, 43.75317, 40 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); RS Cap Sainte Marie, 12.3 km 262° W Marovato, -25.58167, 45.16833, 200 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); RS Cap Sainte Marie, 14.9 km 261° W Marovato, -25.59444, 45.14683, 160 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 3 km E Itampolo, malaise across path of lower bench of Andrimpano Forest, -24.65783, 43.95617, 45 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km E Itampolo, malaise across path of plateau of Andrimpano Forest, -24.65033, 43.96317, 130 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km N Ampotaka, malaise on trail in Vitambany gallery forest, -24.65033, 43.96317, 86 m, Gallery forest (M.E. Irwin, Rin’ha) (CAS); Ambohimahavelona village 33 km NE of Tulear, Andoharano dry forest, -23.44083, 43.89967, 46 m, dry forest (M.E. Irwin, Rin’ha) (CAS); PN Andohahela, Ihazofotsy - Parcel III, transition forest, -24.83483, 46.48683, 80 m, transition between spiny and dry deciduous forests (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Mikea Forest, deciduous dry forest, -22.90367, 43.4755, 30 m, deciduous dry forest (R. Harin’Hala) (CAS); Parcel I, RS Beza Mahafaly, near research station, -23.6865, 44.591, 165 m, dry deciduous forest (R. Harin’Hala) (CAS); PN Tsimanampetsotsa, Mitoho Forest, malaise across trail at escarpment base, -24.0485, 43.75233, 120 m, dense dry forest (M.E. Irwin, Rin’ha) (CAS); Tsimelahy - Parcel II, PN Andohahela, transition forest, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

Camponotus atimo, endemic to Madagascar, generally occurs in the southern part of the island (Fig. 43D). Its habitats range from dry forest and coastal scrub on limestone of the southwestern region to the transitional forest in the extreme southeastern region and Uapaca woodland in the south-central region through the spiny forest and thicket in the extreme south of the island. Field work during the past 25 years has found this species foraging mostly on the ground and through leaf litter, and very rarely on lower vegetation. It usually nests under stones, in rotten logs and soil layers, and rarely in tree stumps.

Figure 43.

Camponotus atimo A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0454042 D distribution map.

Discussion

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%.

Etymology

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

Figs 32A, 33A, 44

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; Wheeler 1922: 1044].

Additional material examined

Mauritius: [Mauritius, 19161], -20.3, 57.583333, (Roger) (CAS); Bassin Blanc, -20.45463, 57.47587, 481 m: (L. Lach) (CAS); Le Pouce, -20.195, 57.522222 (L. Lach) (CAS); Le Pouce, -20.195, 57.522222 (C. M. Courtois) (CAS); Macchabee Forest, -20.4, 57.45, 600 m, disturbed rainforest (P.S. Ward) (MNHN); Petite Rivière Noire Mt. -20.40883, 57.40767, 750 m, rainforest (B.L. Fisher et al.) (UCDC); Petite Rivière Noire Mt., -20.363889, 57.368333 (Michael, Madl) (CAS). Reunion: Mare, Longue, -21.34617, 55.73983, 450 m, rainforest (B.L. Fisher et al.) (ZMHB); Mare Longue, -21.34889, 55.74417, 361 m, Low 1/2wet protected area (J. Casquet) (CAS); Mare Longue, -21.35, 55.75, 150 m, primary forest (F. Blard) (CAS); Mare Longue, -21.363344, 55.753428 (F. Blard) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

The distribution of C. aurosus is restricted to Mauritius and Reunion islands (Fig. 44D). In these islands, this species occupies primary and disturbed rainforest habitats, where workers have been found foraging on the ground and on lower vegetation and nest sites have been located in rotten logs, in the ground, under stones, and in tree trunks.

Figure 44.

Camponotus aurosus A lateral view B head in full-face view C dorsal view of minor worker CASENT0064815 D distribution map.

Discussion

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 becki Santschi, stat. nov.

Figs 27A, 45

Camponotus radamae st. becki Santschi, 1923: 292. Syntype workers, Madagascar (R. Beck) (MHNG) [examined]; 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101992 (MHNG) [examined]. Paralectotype major worker CASENT0101777 (MHNG) [examined]. [First available use of Camponotus (Myrmoturba) maculatus r. radamae var. becki Forel, 1914: 251; unavailable name]. Stat. nov. The worker specimens that are the basis of the unavailable name Camponotus (Myrmoturba) radamae st. becki var altior Santschi, 1923: 292 from Madagascar, Monts Andringitra, point culminant de l’île (Descarpentries) are referred here, AntWeb CASENT0101100 (NHMB) [examined].

Additional material examined

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.) (CAS); PN Andringitra, Plateau d’Andohariana, 39.8 km 204° Ambalavao, -22.18767, 46.90083, 2150 m, rubicole thicket at base of cliff (B.L. Fisher et al.) (CAS); PN Andringitra, Plateau d’Andohariana, base of Pic d’Ivangomena, -22.2, 46.9, 2050 m, montane rainforest, (Goodman leg.) (CAS); PN Andringitra, 8.5 km SE Antanitotsy, -22.16667, 46.96667, 1990 m, montane rainforest (Sylvain) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

Based on recent collection data, Camponotus becki is restricted to the ericoid thickets and montane rainforests of the PN Andringitra (Fig. 45 D). The species has been found nesting under stones and foraging on the ground.

Figure 45.

Camponotus becki A lateral view B head in full-face view C dorsal view of minor worker CASENT0071380 D distribution map.

Discussion

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.

Camponotus bemaheva sp. nov.

http://zoobank.org/1AF0EE48-7F21-4744-961F-86BCEE7A9029

Figs 7B, 13A, 46

Holotype worker

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 (CAS).

Additional material examined

Madagascar: Antsiranana: RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (CAS). Fianarantsoa: Forêt d’Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CAS). Mahajanga: PN Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) (CAS). Toliara: 48 km ENE Morondava, -20.06667, 44.65, 30 m, tropical dry forest (D.M. Olson) (PSWC), Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush (Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Betioky ; RS Beza Mahafaly, Parcelle Belle vue, 07 km W of Research Station, -23.68983, 44.5755, 177 m, spiny forest, (Rin’ha) (CAS); Atsimo Andrefana Region, District of Betioky, 30 km E Betioky, RS Beza Mahafaly (Around Research Station), -23.6865, 44.591, 165 m, Galery dry deciduous forest (Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Sakaraha, PN Zombitse; 60 m E of ANGAP Entrance office, -22.8865, 44.69217, 838 m, deciduous spiny forest (Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Sakaraha, PN Zombitse; 900 m N from ANGAP Entrance office, -22.8405, 44.73117, 823 m, spiny deciduous forest (Rin’Ha, Mike) (CAS); FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); Forét de Kirindy, -20.0671, 44.65723, 50 m (H. Wood & J. Miller) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.06915, 44.66042, 30 m, tropical dry forest (B.L. Fisher et al.) (CAS); Makay Mts., -21.227, 45.33222, 475 m, Gallery forest on sandy soil (B.L. Fisher et al.) (CAS); Manombo, -22.80707, 43.76375, 222 m, gallery forest, TS1 (Frontier Wilderness Project) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza dry forest 07,5 Km E of Beroboka, -19.9775, 44.66633, 50 m, dry forest (M. Irwin, Rin’ha) (CAS); Menabe Region, District of Morondava, Beroboka village, 45 km NE of Morondava, Antsarongaza galery forest, 07 km E of Beroboka, -19.9775, 44.66533, 45 m, Galery forest (M. Irwin, Rin’ha) (CAS); PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Vohibasia Forest, 59 km NE Sakaraha, -22.46667, 44.85, 780 m (Sylvain) (CAS); near ANGAP office, PN Zombitse, -22.8865, 44.69217, 840 m, deciduous spiny forest (R. Harin’Hala) (CAS); Parcel I, RS Beza Mahafaly, near research station, -23.6865, 44.591, 165 m, dry deciduous forest (R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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

Distribution and biology

Camponotus bemaheva is restricted to the spiny bush and thicket of the southern region and the dry forest habitats of western Madagascar (Fig. 46D). The few colony nests that have been found were collected from rotten logs and under stones, but workers have been collected while foraging on the ground or on low vegetation.

Figure 46.

Camponotus bemaheva A lateral view B head in full-face view C dorsal view of minor worker CASENT0154158 D distribution map.

Discussion

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.

Etymology

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 boivini Forel, stat. rev.

Figs 23B, 47

Camponotus maculatus r. boivini Forel, 1891: 34. Syntype workers, queen and male, Madagascar (Sikora) (MHNG); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101343 (MHNG) [examined]. Paralectotypes with same data as lectotype but: 1 minor worker CASENT0101349, 2 major workers CASENT0101342, CASENT0101348 and 1 male CASENT0101607 (MHNG) [examined]. Raised to species by Dalla Torre 1893: 223. As subspecies of Camponotus maculatus by Emery, 1895: 337; Wheeler 1922: 1040. As subspecies of Camponotus hova by Emery, 1920b: 6, 1925: 86. Stat. rev.

Camponotus maculatus st. fairmairei Santschi, 1911a: 130. Syntype workers Madagascar, colony living with larvae of Fulgorids (Fairmaire, 1900) (NHMB); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101102 (NHMB) [examined]. [Camponotus hova fairmairei: as subspecies of Camponotus hova by Emery, 1920b: 6; Emery 1925: 86]. Syn. nov.

Note

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.

Additional material examined

Madagascar: Antananarivo: Alasora, -18.96245, 47.58925, 1434 m, eucalyptus plantation (B.L. Fisher et al.) (CAS); Andohony I Non Protected Area, 22.62 km SW Antsirabe, -20.06784, 46.99068, 1451 m, Savannah grassland (A. Ravelomanana) (CAS); Andohony II Non Protected Area, 22.71 km SW Antsirabe, -20.06904, 46.99199, 1398 m, Savannah grassland (A. Ravelomanana) (CAS); Ankalalahana, -18.99666, 47.1118, 1353 m, Uapaca woodland, (H.E. Marti et al.) (CAS); Ankalalahana, -19.00659, 47.1122, 1375 m, Uapaca woodland (B.L. Fisher et al.) (CAS); Antaponimanadala III Non Protected Area, 6.55 km E Manalalondo, -19.25583, 47.17751, 1987 m, Savannah grassland (A. Ravelomanana) (CAS); Antaponimanadala IV Non Protected Area, 6.66 km E Manalalondo, -19.25361, 47.17634, 1951 m, Savannah grassland (A. Ravelomanana) (CAS); Antsirabe, -19.866, 47.0355, 1550 m, urban/garden (B.L. Fisher et al.) (CAS); Beapombo III Non Protected Area, 22.30 km SW Antsirabe, -20.0662, 46.99839, 1612 m, Uapaca woodland (A. Ravelomanana) (CAS). Antsiranana: Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CAS); Forêt Ambato, 26.6 km 33° Ambanja, -13.4645, 48.55167, 150 m, rainforest (B.L. Fisher) (CAS); Forêt d’Andavakoera, 21.4 km 75° ENE Ambilobe; 4.6 km 356° N Betsiaka, -13.11833, 49.23, 425 m, rainforest (B.L. Fisher) (CAS); Forêt d’Ampondrabe, 26.3 km 10° NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest (B.L. Fisher) (CAS); Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) (CAS); Forêt d’Orangea, 3.6 km 128° SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) (CAS); Galoko chain, Mont Galoko, -13.58745, 48.71419, 380 m, rainforest (B.L. Fisher et al.) (CAS); Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest, (J.-J. Rafanomezantsoa et al.) al. (CAS); Nosy Ankao forest, -12.79164, 49.82378, 25 m, tropical dry forest (B.L. Fisher et al.) (CAS); Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CAS); Réserve Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) (CAS); Réserve Analamerana, 28.4 km 99° Anivorano-Nord, -12.74667, 49.49483, 60 m, tropical dry forest (B.L. Fisher) (CAS); RS Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (Fisher, Griswold et al.) (CAS); RS Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest (Fisher, Griswold et al.) (CAS); RS Ankarana, Anilotra, -12.90981, 49.11057, 98 m, Dry forest on tsingy (B.L. Fisher et al.) (CAS). Fianarantsoa: 28 km SSW Ambositra, Ankazomivady, -20.775, 47.16833, 1670 m, grassland (B.L. Fisher) (CAS); Amoron’i Mania Region, District of Ambositra, Italaviana Uapaca forest, 35 km SE of Antsirabe, -20.17333, 47.086, 1359 m, Uapaca forest (Rin’Ha, Mike) (CAS); PN Isalo, Ampangabe I Non Protected Area, 21.4 km W Itremo, -20.61111, 46.60688, 1414 m, savannah woodland (A. Ravelomanana) (CAS); Ampangabe IV Non Protected Area, 21.37 km W Itremo, -20.61278, 46.60774, 1417 m, savannah woodland (A. Ravelomanana) (CAS); Antohatsahomby IV Non Protected Area, 22.67 km NW Itremo, -20.56306, 46.58097, 1708 m, Uapaca woodland (A. Ravelomanana) (CAS); Antohatsahomby V Non Protected Area, 22.63 km NW Itremo, -20.56722, 46.57923, 1726 m, Uapaca woodland (A. Ravelomanana) (CAS); Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, grassland (Fisher, Griswold et al.) (CAS); Manandriana II Non Protected Area, 27.12 km SW Ambositra, -20.7325, 47.09461, 1589 m, Savannah grassland (A. Ravelomanana) (CAS); Soanierenana II Non Protected Area, 25.61 km SW Ambositra, -20.72194, 47.10896, 1732 m, savannah grassland (A. Ravelomanana) (CAS); Soanierenana IV Non Protected Area, 25.22 km SW Ambositra, -20.72389, 47.10705, 1736 m, savannah grassland (A. Ravelomanana) (CAS); Soanierenana IV Non Protected Area, 25.22 km SW Ambositra, -20.72389, 47.10705, 1736 m, savannah grassland (A. Ravelomanana) (CAS). Mahajanga: Boeny Region, District of Marovoay, PN Ankarafantsika, Ampijoroa SF, 160 km North of Maevatanana on RN 04, -16.31933, 46.81333, 42 m, deciduous forest (Mike, Rin’ha) (CAS); Boeny Region: PN Ankarafantsika, Ambodimanga, -16.32342, 46.82443, 75 m, dry forest, Canopy Bootcamp, (A. Karunakaran) (CAS). Forêt Ambohimanga, 26.1 km 314° Mampikony, -15.96267, 47.43817, 250 m, tropical dry forest (B.L. Fisher) (CAS). Forêt de Tsimembo, 11.0 km 346° NNW Soatana, -18.99528, 44.4435, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS). Mahavavy River, 6.2 km 145° SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) (CAS). Manerinerina, 76.6 km N Antsohihy, -14.10744, 48.11046, 247 m, disturbed forest (B.L. Fisher et al.) (CAS). PN Ankarafantsika, Ampijoroa SF, 5.4 km 331° NW Andranofasika, -16.29889, 46.813, 70 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Ankarafantsika, Ampijoroa SF, 40 km 306° NW Andranofasika, -16.32083, 46.81067, 130 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Baie de Baly, 12.4 km 337° NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 9.8 km 300° WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CAS); PN Tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River, -19.13222, 44.81467, 100 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CAS); PN Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Réserve d’Ankoririka, 10.6 km 13° NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Réserve forestière Beanka, 50.7 km E Maintirano, -17.88021, 44.46877, 140 m, tropical dry forest edge (B.L. Fisher et al.) (CAS); Réserve forestière Beanka, 52.7 km E Maintirano, -18.0622, 44.52587, 300 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CAS); Réserve forestière Beanka, 53.6 km E Maintirano, -18.04014, 44.53394, 272 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CAS); RS Bemarivo, 23.8 km 223° SW Besalampy, -16.925, 44.36833, 30 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Sofia Region, District of Port-Berger, Ambovomamy 20 km N of Port-Berger, -15.45117, 47.61333, 86 m, secondary forest on white sandy area (Mike, Frank, Rin’ha) (CAS). Toamasina: Ambatovy, 12.4 km NE Moramanga, -18.85813, 48.28488, 1040 m, grassland (B.L. Fisher et al.) (CAS); PN Andasibe, botanic garden near entrance, west of ANGAP office, -18.925172, 48.418651, 1025 m, tropical forest (M.E. Irwin, R. Harin’Hala) (CAS). Toliara: Forêt de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.06855, 44.65956667, 30 m, tropical dry forest (B.L. Fisher) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.045, 44.66222, 100 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Tsinjoriaky, 6.2 km 84° E Tsifota, -22.80222, 43.42067, 70 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Makay Mts., -21.30997, 45.12946, 590 m, dry forest on sandy soil (B.L. Fisher et al.) (CAS); Makay Mts., -21.31664, 45.1296, 620 m, dry forest on sandy soil (B.L. Fisher et al.) (CAS); Makay Mts., -21.25864, 45.16412, 500 m, gallery forest with bamboo (B.L. Fisher et al.) (CAS); Makay Mts., -21.21985, 45.32396, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) (CAS); Makay Mts., -21.20978, 45.34184, 525 m, gallery forest on sandy soil (B.L. Fisher et al.) (CAS); PN Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, -23.99222, 43.88067, 90 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 5 km E Itampolo, Malaise across path of plateau of Andrimpano Forest, -24.65033, 43.96317, 130 m, dry forest (M.E. Irwin, Rin’ha) (CAS); RS Kalambatritra, Ambinanitelo, -23.45373, 46.45773, 1345 m, grassland (B.L. Fisher et al.) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

The distribution of C. boivini is generally limited to western and the high plateau of Madagascar (Fig. 47D). This species occurs mostly in dry forest habitats in the north through spiny forest areas in the south. Along its north-south range it can be found in gallery forests, disjunct montane rainforest, Uapaca woodlands, and savannah grasslands. When nesting in rotten logs, rotting tree stumps, under stones, in dead branches, twigs on the ground, and in the soil, C. boivini typically forages in leaf litter and seldom on the forest floor and low vegetation.

Figure 47.

Camponotus boivini A lateral view B head in full-face view C dorsal view of minor worker CASENT0486575 D distribution map.

Discussion

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.

Santschi (1911a) originally described Camponotus maculatus st. fairmairei without any comparison of the specimens to those of other species, though his description corresponds to the type specimens of C. boivini. The observation of the samples and the distributional data obtained from the recent ant survey in Madagascar combined with this information are sufficiently enough to reasonably synonymize Camponotus maculatus st. fairmairei here.

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%.

Camponotus bozaka sp. nov.

http://zoobank.org/CAD5FB8A-8FF4-47C6-9590-A7E8342C6839

Figs 26A, 28B, 48

Holotype worker

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 (CAS).

Paratype

1 major worker with same data as holotype but with specimen code CASENT0071370 (CAS).

Additional material examined

Madagascar: Fianarantsoa: Manandriana I Non Protected Area, 27.11 km SW Ambositra, -20.73194, 47.09413, 1590 m, Savannah grassland (A. Ravelomanana) (CAS); Manandriana III Non Protected Area, 27.25 km SW Ambositra, -20.73333, 47.09391, 1578 m, Savannah grassland (A. Ravelomanana) (CAS); PN Andringitra, Plateau d’Andohariana, 39.8 km 204° Ambalavao, -22.18767, 46.90083, 2150 m, rubicole thicket at base of cliff (B.L. Fisher et al.) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

Known only from the south-central high plateau of Madagascar, C. bozaka occupies rubicole thicket, Uapaca woodland, savannah grassland, and shrubland habitats (Fig. 48D). Worker specimens have been found foraging through leaf litter and on low vegetation. The nest sites are in the ground or under stones.

Figure 48.

Camponotus bozaka A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0217309 D distribution map.

Discussion

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.

Etymology

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 cemeryi Özdikmen, stat. nov.

Figs 21A, 49

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 (NHMB) [examined]. Paralectotypes: 2 major workers with the same data as lectotype but with specimen codes CASENT0101104 (NHMB), CASENT0101894 (MHNG); 1 alate queen CASENT0101848 and 1 male CASENT0101911 (MHNG) [examined]. Replacement name of Camponotus hova var. luteolus Emery, 1925: 85. Stat. nov.

Note

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].

Additional material examined

Madagascar: Antananarivo: Navoatra VI Non Protected Area, 7.3 km NW Arivonimamo, -18.97889, 47.12253, 1276 m, Uapaca woodland (A. Ravelomanana) (CAS). Antsiranana: Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) (CAS). Fianarantsoa: 28 km SSW Ambositra, Ankazomivady, -20.775, 47.16833, 1670 m, grassland (B.L. Fisher) (CAS); Ambalavao, -21.83267, 46.93867, 1020 m, urban/garden (B.L. Fisher et al.) (CAS); Ampandravelo III Non Protected Area, 10.72 km NE Ranohira, -22.53944, 45.51497, 869 m, Shrubland (A. Ravelomanana) (CAS); Ampotoampoto I National Parc, 8.02 km NW Ilakaka, -22.62833, 45.18859, 917 m, savannah woodland (A. Ravelomanana) (CAS); Ampotoampoto IV National Parc, 7.83 km NW Ilakaka, -22.62944, 45.1912, 923 m, savannah woodland (A. Ravelomanana) (CAS); dry wash, 1 km E of PN Isalo Interpretive Center, -22.62667, 45.35817, 885 m, dry wash (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Forêt d’Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Horombe Region, District of Ihosy, PN Isalo, 900 m E of ANGAP Interpretation Center, -22.62667, 45.35817, 701 m, open area near stream (Rin’Ha, Mike) (CAS); Horombe Region, Ihosy Distric, PN Isalo, 1 km E of ANGAP Interpretation Center, -22.62667, 45.35817, 823 m, open area (Rin’Ha, Mike) (CAS); Isalo II National Parc, 12.26 km SW Ranohira, -22.61528, 45.31307, 867 m, Bismarckia woodland (A. Ravelomanana) (CAS); Isalo III National Parc, 12.02 km SW Ranohira, -22.61583, 45.31084, 870 m, Bismarckia woodland (A. Ravelomanana) (CAS); Isalo IV National Parc, 12 km SW Ranohira, -22.61472, 45.31304, 867 m, Bismarckia woodland (A. Ravelomanana) (CAS); RS Manombo, 32 km SE of Farafangana, -23.02183, 47.72, 36 m, Lowland rainforest (Rin’Ha, Mike) (CAS); PN Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland (Fisher, Griswold et al.) (CAS); PN Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CAS); radio tower, PN Ranomafana, -21.25833, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); stream area, 900 m E of PN Isalo Interpretive Center, -22.62667, 45.35817, 750 m, open area near stream (R. Harin’Hala) (CAS). Mahajanga: PN Ankarafantsika, Ampijoroa SF, 160 km N Maevatanana, deciduous forest, -16.31944, 46.81333, 43 m, deciduous forest, Mike (Irwin, Rin’ha), Harin’Hala (CAS); Boeny Region, District of Soalala, Beaboaly Bamboo forest Station10 km SW of Soalala, 04 km of Baly village, -16.04533, 48.804, 9 m, Bamboo Forêt (Mike, Rin’ha) (CAS); Boeny Region, District of Marovoay, PN Ankarafantsika, Ampijoroa SF, 160 km North of Maevatanana on RN 04, -16.31933, 46.81333, 42 m, deciduous forest (Mike, Rin’ha) (CAS); Boeny Region, District of Soalala Analamanitra forest, 14 km SW of Mitsinjo, -16.7, 45.7, 19 m, dense dry forest (Mike, Rin’ha) (CAS); Mahavavy River, 6.2 km 145° SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) (CAS); Melaky Region, District of Mintirano, Ampasy 50 km E of Maintirano, -18.004, 44.452, 85 m, dry forest (Mike, Rin’ha) (CAS); PN Baie de Baly, 12.4 km 337° NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 9.8 km 300° WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Réserve forestière Beanka, 48.9 km E Maintirano, -18.02472, 44.48788, 250 m, savannah shrubland (B.L. Fisher et al.) (CAS); Réserve forestière Beanka, 50.2 km E Maintirano, -18.02127, 44.49566, 250 m, savannah woodland (B.L. Fisher et al.) (CAS); Réserve forestière Beanka, 50.7 km E Maintirano, -17.8873, 44.47113, 160 m, savannah shrubland (B.L. Fisher et al.) (CAS). Toliara: Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush (Rin’Ha, Mike) (CAS); Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush (Rin’Ha, Mike) (CAS); Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 152 m, Bush (Mike, Rin’ha) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.00667, 46.30333, 85 m, Galery forest (Rin’Ha, Mike) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.00667, 46.30333, 85 m, Galery forest (Rin’Ha, Mike) (CAS); Anosy Region, District of Amboasary, PN Andohahela, Parcelle III, Ihazofotsy, 32 km NE Amboasary, -24.83083, 46.53617, 58 m, dry forest, spiny forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.021, 46.3055, 36 m, spiny forest (Mike, Rin’ha) (CAS); Anosy Region, District of Fort-Dauphin, PN Andohahela, Parcelle II, Tsimela, 42 km W of Fort-Dauphin, -24.93683, 46.62667, 176 m, transition forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82505, 46.57811, 90 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82466, 46.60111, 100 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo Andrefana Region, District of Betioky, 30 km E Betioky, RS Beza Mahafaly (Around Research Station), -23.6865, 44.591, 165 m, Galery dry deciduous forest (Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Tulear II, Mikea deciduous dry forest 3 km N Andranomavo village, -22.90367, 43.4755, 30 m, Deciduous dry forest (Rin’Ha, Mike) (CAS); Beza-Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CAS); Forêt de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.045, 44.66222, 100 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Tsinjoriaky, 6.2 km 84° E Tsifota, -22.80222, 43.42067, 70 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Makay Mts., -21.29961, 45.12919, 570 m, Dry forest edge and burned savannah (B.L. Fisher et al.) (CAS); Makay Mts., -21.34109, 45.18054, 500 m, Barren rock with sparse vegetation, burned grass (B.L. Fisher et al.) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza galery forest 07 km E of Beroboka, -19.9775, 44.66533, 45 m, Galery forest (M. Irwin, Rin’ha) (CAS); PN Andohahela, Forêt d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, -24.93, 46.6455, 300 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Kirindy Mite, 16.3 km 127° SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, -23.99222, 43.88067, 90 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Mitoho Cave, 6.4 km 77° ENE Efoetse, 17.4 km 170° S Beheloka, -24.04722, 43.75317, 40 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CAS); RS Beza Mahafaly, Parcel 1, -23.65, 44.63333, 130 m, tropical dry forest (P.S. Ward) (CAS); southern Isoky-Vohimena Forest, 59 km NE Sakaraha, -22.46667, 44.85, 730 m, tropical dry forest (B.L. Fisher) (CAS); 3 km E Itampolo, malaise across path of lower bench of Andrimpano Forest, -24.65783, 43.95617, 45 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km E Itampolo, malaise across path of plateau of Andrimpano Forest, -24.65033, 43.96317, 130 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km N Ampotaka, malaise on trail in Vitambany gallery forest, -24.65033, 43.96317, 86 m, Gallery forest (M.E. Irwin, Rin’ha) (CAS); Berenty Special Reserve, 8 km NW Amboasary, 58 km SW of Fort Dauphin, -25.00667, 46.30333, 85 m, gallery forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); PN Andohahela, Ihazofotsy - Parcel III, transition forest, -24.83483, 46.48683, 80 m, tropical dry forest, transition between spiny and dry deciduous forests (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Mikea Forest, deciduous dry forest, -22.90367, 43.4755, 30 m, deciduous dry forest (R. Harin’Hala) (CAS); Mikea Forest, spiny forest, -22.91333, 43.48222, 37 m, spiny forest (R. Harin’Hala) (CAS); near ANGAP office, PN Zombitse, -22.8865, 44.69217, 840 m, deciduous spiny forest (R. Harin’Hala) (CAS); Parcel I, RS Beza Mahafaly, near research station, -23.6865, 44.591, 165 m, dry deciduous forest (R. Harin’Hala) (CAS); Parcel II, RS Beza Mahafaly, near Bellevue, -23.68983, 44.5755, 180 m, spiny forest (R. Harin’Hala) (CAS); PN Tsimanampetsotsa, Mitoho Forest, malaise on plateau, -24.0485, 43.75233, 150 m, dense dry forest (M.E. Irwin, Rin’ha) (CAS); Tsimelahy - Parcel II, PN Andohahela, transition forest, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

Camponotus cemeryi occurs in dry forest and spiny forest of west Madagascar and in the savannah shrubland and woodland of the central plateau (Fig. 49D). Along its distribution, members of the species can also be found in gallery forest habitats and in transitional forest between dry and humid forests. It may also establish colonies in human-modified habitats on the high plateau. Specimens have been collected mostly foraging on the forest floor and rarely on lower vegetation. The species nests in rotten logs, in the ground, under stones, and in rotting tree stumps.

Figure 49.

Camponotus cemeryi A lateral view B head in full-face view C dorsal view of minor worker CASENT0217306 D distribution map.

Discussion

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

Figs 8A, 9B, 10B, 50

Camponotus cervicalis Roger, 1863: 134. Syntype workers, Madagascar (Humboldt) (NHMB); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101178 (NHMB) [examined]. Paralectotypes: 3 minor workers and 2 major workers of same data as lectotype but respectively specimen coded as: CASENT0101551, CASENT0101552 (MHNG), CASENT0104634 (ZMHB); CASENT0101704 and CASENT0101779 (MHNG) [examined]. Combination in Camponotus (Dinomyrmex): Forel, 1914: 268; in Camponotus (Tanaemyrmex): Emery, 1925: 84.

Camponotus gaullei Santschi, 1911a: 128. Syntype workers, Madagascar, S. de la Baie d’Antongil, (NHMB); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101179 (NHMB) [examined]. [Combination in Camponotus (Dinomyrmex): Wheeler, 1922: 1043; in Camponotus (Tanaemyrmex): Emery, 1925: 84]. As subspecies of Camponotus cervicalis by Emery, 1925: 84. Syn. nov.

Camponotus perroti Forel, 1897: 202. Syntype workers, Madagascar, Sainte Marie (Perrot) (MHNG); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101350 (MHNG) [examined]. Paralectotype major worker of same data as lectotype but with specimen code CASENT0101351 (MHNG) [examined]. Syn. nov.

Camponotus perroti aeschylus Forel, 1913: 224. Holotype (by monotypy) alate queen, Madagascar (C. Keller) AntWeb CASENT0101561 (MHNG) [examined]. Syn. nov.

Camponotus gerberti Donisthorpe, 1949: 271. Syntype workers, Madagascar, Sainte Marie (R. Géberti 1917) (NHMUK); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102310 (NHMUK) [examined]. Paralectotypes: 2 major workers of the same data as lectotype but with the following specimen codes: CASENT0102309, CASENT0102311 (NHMUK) [examined]. [Combination in Camponotus (Tanaemyrmex): Donisthorpe, 1949: 271]. Syn. nov.

Additional material examined

Madagascar: Antananarivo: [Madagascar]; Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (Humblot) (MNHN); Iharanandriana, -19.15823, 47.49702, 1513 m, Uapaca woodland (B.L. Fisher et al.) (CAS). Antsiranana: Forêt Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CAS); RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, rainforest (P.S. Ward) (PSWC). Toamasina: S. de la baie d’Antongil. (NHMB); Mananara Avaratra, -16.170555, 49.765208, 18 m (B.L. Fisher et al.) (CAS); Nosy Mangabe, -15.5, 49.76667, <5 m, rainforest edge (P.S. Ward) (PSWC); Nosy Mangabe, 7.43 km S Maroantsetra, -15.4973, 49.76223, 3 m, littoral rainforest (B.L. Fisher et al.) (CAS); PN Masoala, 39.4 km 150° SSE Maroantsetra, -15.71, 49.97, 200 m, rainforest (B.L. Fisher & H.J. Ratsirarson) (CAS); Parcelle K7 Tampolo, -17.28333, 49.41667, 10 m, littoral forest (Malagasy ant team) (CAS); RNI Betampona, 34.1 km 332° Toamasina, -17.91614, 49.20185, 550 m, rainforest (B.L. Fisher et al.) (CAS); SF Tampolo, 10 km NNE Fenoarivo Atn. -17.2825, 49.43, 10 m, littoral rainforest (B.L. Fisher) (CAS); Tampolo, -17.28333, 49.41667, 10 m, littoral forest (Malagasy ant team) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 50D). Workers of the species have been found foraging on the ground and through leaf litter, but nest sites range from rotten logs, rotting tree stumps, rot pockets above the ground, live trees, under tree bark, and in moss and leaf litter.

Figure 50.

Camponotus cervicalis A lateral view B head in full-face view C dorsal view of minor worker CASENT0217303 D distribution map.

Discussion

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, Santschi (1911a) discussed the resemblance of the species to C. cervicalis and C. dufouri and no strong separating characters were provided. However, with recent extensive sampling of the Malagasy ant fauna, the observation of the range of worker class sizes in these latter species indicates the closer similarity between C. cervicalis and C. gaullei. Combined with the close distance between the type localities of both species, this information would be enough to synonymize C. gaullei under C. cervicalis.

The description of C. perroti by Forel (1897) was based on a limited number of worker specimens which had a close similarity to C. cervicalis and C. dufouri. Careful examinations of more specimens obtained from the recent ant surveys in Madagascar show that C. perroti and C. cervicalis belong to the same species and that C. perroti merits its placement as a junior synonym of C. cervicalis.

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%.

Camponotus daraina sp. nov.

http://zoobank.org/10EF7582-8BEC-438F-A757-BE96FF472CDB

Figs 14A, 51

Holotype worker

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 (CAS).

Paratypes

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 (NHMUK, MHNG, MSNG, PBZT, CAS).

Additional material examined

Madagascar: Antsiranana: Forêt Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CAS); Forêt d’Antsahabe, 11.4 km 275° W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher et al.) (CAS); Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 650–800 m, rainforest (B.L. Fisher et al.) (CAS); RS Manongarivo, 12.8 km 228° SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CAS); PN Montagne d’Ambre [1^st campsite], 960 m, rainforest (R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 51D). Camponotus daraina is strictly terrestrial; its workers forage mainly on the forest floor and through leaf litter, while its nests have been found in rotten logs and under stones.

Figure 51.

Camponotus daraina A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0077433 D distribution map.

Discussion

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%.

Etymology

The species name daraina is a singular non-Latin noun used in apposition and refers to the type locality.

Camponotus dufouri Forel

Figs 8B, 12A, 52

Camponotus dufouri Forel, 1891: 16, 74 (Key). Syntype workers, Madagascar, 30 miles NW Tamatave (O’Swald) (MHNG) [examined]; 1 syntype major worker designated as lectotype, by present designation, Madagascar, AntWeb CASENT0101644 (MHNG) [examined]. Paralectoype: 1 major worker same data as lectotype but with specimen code: CASENT0101669 (MHNG) [examined]. [Combination in Camponotus (Dinomyrmex): Forel, 1914: 268; in Camponotus (Tanaemyrmex): Emery, 1925: 85].

Camponotus dufouri imerinensis Forel, 1891: 18. Syntype workers and queen, Madagascar, Environ d’Antananarivo, Imerina [labeled as “Andrangoloaka”] (Sikora) (MHNG) [examined]; 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101812 (Sikora) (MHNG) [examined]. Paralectotypes: 1 major worker and 1 alate queen of the same data as lectotype but with the following specimen codes: CASENT0101919 (Sikora) and CASENT0101679 (Camboué) (MHNG) [examined]. Syn. nov.

Additional material examined

Madagascar: Antananarivo: [Madagascar], Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (Sicora) (CAS); 3 km 41 °NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (Fisher, Griswold et al.) (CAS); Région Analamanga, SF Mandraka, -18.9183, 47.91687, 1285 m, montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); [Andrangoloaca]; Mantasoa; Manjakandriana, -19.033333, 47.9166666, 1409 m (MHNG). Antsiranana: 9.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.46667, 1280 m, montane rainforest (B.L. Fisher) (CAS); 9.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) (CAS); Binara Forest, -13.26392, 49.59919, 1065 m, rainforest (B.L. Fisher et al.) (CAS); Binara Forest, -13.26388, 49.60141, 900 m, rainforest (B.L. Fisher et al.) (CAS); Forêt Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CAS); Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 650–800 m, rainforest (B.L. Fisher) (CAS); Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 800 m, rainforest (B.L. Fisher) (CAS); Makirovana forest, -14.16044, 49.95216, 550 m, rainforest (B.L. Fisher et al.) (CAS); PN Masoala, -15.32331, 50.30751, 60 m, rainforest (B.L. Fisher et al.) (CAS); PN Montagne d’Ambre, Roussette Camp 7 km SW Park entrance, -12.51444, 49.18139, 960 m, rainforest (Mike, Frank, Rin’ha) (CAS); PN Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, -14.44333, 49.74333, 1325 m, montane rainforest (B.L. Fisher) (CAS); PN Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327 °NNW Manantenina, -14.435, 49.76, 775 m, rainforest (B.L. Fisher et al.) (CAS); PN Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333 °NNW Manantenina, -14.43667, 49.775, 450 m, rainforest (B.L. Fisher) (CAS); PN Montagne d’Ambre, 3.6 km 235° SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CAS); PN Montagne d’Ambre, Antomboka, -12.51269, 49.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d’Ambre, Crête, -12.58132, 49.13368, 1110 m, montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d’Ambre, Petit lac, -12.53664, 49.17412, 1130 m, montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d’Ambre, Roussettes, -12.52574, 49.17238, 1025 m, montane rainforest (B.L. Fisher et al.) (CAS); R.N.I. Marojejy, 11 km NW Manantenina, -14.43333, 49.75, 1225 m, montane rainforest (E.L. Quinter) (CAS); RS Manongarivo, 10.8 km 229 °SW Antanambao, -13.96167, 48.43333, 400 m, rainforest (B.L. Fisher) (CAS); RS Manongarivo, 12.8 km 228° SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CAS); RS Manongarivo, 14.5 km 220° SW Antanambao, -13.99833, 48.42833, 1175 m, montane rainforest (B.L. Fisher) (CAS); PN Marojejy, 10 km NW Manantenina, -14.43333, 49.76667, 750 m, rainforest (E.L. Quinter) (CAS); PN Marojejy, 8 km NW Manantenina, -14.43333, 49.78333, 450 m, rainforest (E.L. Quinter) (CAS); Sakalava Beach [vegetated beach dunes], -12.26972, 49.39167, 10 m, across sandy trail in dwarf litoral forest (R. Harin’Hala) (CAS); SAVA Region, District of Sambava, PN Marojejy, 5 km W of Manantenina village, 1^st Camp site (Mantella), -14.43817, 49.774, 487 m, Low altitude rainforest (Rin’Ha, Mike) (CAS); Montaigne Francais, 150 m, along forested limestone ridge (R. Harin’Hala) (CAS); Montaigne Francais, 150 m, along forested limestone ridge (R. Harin’Hala) (CAS); PN Montagne d’Ambre [1^st campsite], 960 m, rainforest (Irwin, Schlinger, Harin’H) (CAS); PN Montagne d’Ambre [lemur trail], 975 m, rainforest (R. Harin’Hala) (CAS); PN Montagne d’Ambre [Petit Lac road], -12.533333, 49.166667, 1125 m, rainforest (Irwin, Schlinger, Harin’H) (CAS). Fianarantsoa: 40 km S Ambalavao, PN Andringitra, -22.21667, 46.96667, 1275 m, montane rainforest (B.L. Fisher) (CAS); 45 km S Ambalavao, -22.21667, 47.01667, 720 m, rainforest edge (B.L. Fisher) (MHNG); Andrambovato along river Tatamaly, -21.51082, 47.40992, 1063 m, rainforest (B.L. Fisher et al.) (CAS); Belle Vue trail, PN Ranomafana, -21.2665, 47.42017, 1020 m, mixed tropical forest (M.E. Irwin, F.D. Parker (R. Harin’Hala) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana Belle vue area1.2 km S of PN Ranomafana entrance, -21.2665, 47.42017, 1018 m, rainforest (Rin’Ha, Mike) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana, 12 km W of Ranomafana, -21.25083, 47.40717, 1127 m, forest edge, open area (Rin’Ha, Mike) (CAS); Forêt d’Ambalagoavy Nord, Ikongo, Ambatombe, -21.857068, 47.37849, 625 m (R. Harin’Hala & M.E. Irwin) (CAS); Forêt de Vevembe, 66.6 km 293° Farafangana, -22.791, 47.18183, 600 m, rainforest, transition to montane forest (B.L. Fisher et al.) (CAS); JIRAMA water works near river, PN Ranomafana, -21.2485, 47.45217, 690 m, open area near stream (R. Harin’Hala) (CAS); PN Ranomafana; Ranomafana; Ifanadiana, -21.2161494, 47.4565003, 1087 m (B. Pettersson) (CAS); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); PN Ranomafana, Talatakely, -21.24833, 47.42667 (CE Griswold, DH Kavanaugh, ND Penny, MJ Raherilalao, JS Ranorianarisoa, J Schweickert) (CAS); RS Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (B.L. Fisher, Sylvain) (CAS); RS Ivohibe, 7.5 km ENE Ivohibe, -22.47, 46.96, 900 m, rainforest (B.L. Fisher, Sylvain) (CAS); radio tower, PN Ranomafana, -21.25833, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M. Irwin, R. Harin’Hala) (CAS); PN Ranomafana, Talatakely area, 0.4 km WSW of Park Entrance, -21.41667, 47.68333, 900 m, mixed tropical forest (D.H. & K.M. Kavanaugh) (CAS); Réserve Speciale Manombo 24.5 km 228° Farafangana, -23.01583, 47.719, 30 m, rainforest (B.L. Fisher et al.) (CAS); Vohiparara broken bridge, -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin’Hala) (CAS). Toamasina: [Bois 30 miles NW. Tamatave], Ambodiriana; Toamasina Rural, -17.88832, 49.22849, 373 m (Swald) (CAS); [Tamatave?], Andranobolahy, Toamasina Rural, -18, 49, 370 m (H. O’Swald) (CAS); 5.3 km SSE Ambanizana, Andranobe, -15.66667, 49.96667, 600 m, rainforest (B.L. Fisher) (MHNG); 5.3 km SSE Ambanizana, Andranobe, -15.67133, 49.97395, 425 m, rainforest (B.L. Fisher) (MHNG); 6.2 km SSE Ambanizana, Be Dinta, -15.66667, 49.99806, 600 m, rainforest (V. Razafimahatratra.) (MNHN); 6.3 km S Ambanizana, Andranobe, -15.6813, 49.958, 25 m, rainforest (B.L. Fisher) (MNHN); 6.9 km NE Ambanizana, Ambohitsitondroina, -15.58506, 50.00952, 825 m, rainforest (B.L. Fisher) (MNHN); 7 km SE PN Andasibe Headquarters, -18.969856, 48.465894, 1050 m, tropical forest (M.E. Irwin R. Harin’Hala) (CAS); Analanjirofo Region, District of ToamasinA Mobot Site, Analalava humid dense forest low altitude on the sand 7 km SW of Foulpointe, -17.69333, 49.46028, 75 m, dense humide low alt on the sandy soil (Mike, Rin’ha) (CAS); PN Andasibe, botanic garden near entrance, West of ANGAP office, -18.925172, 48.418651, 1025 m, tropical forest (M.E. Irwin R. Harin’Hala) (CAS); Ankerana, -18.40829, 48.82107, 750 m, rainforest (B.L. Fisher et al.) (CAS); Antsianaka; Ambatosoratra; Ambatondrazaka, -17.58333, 48.5, 752 m (Perrot Freres) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80398, 48.40358, 1064 m, rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.79944, 48.40375, 1016 m, rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Tsaravoniana, -18.76124, 48.42134, 939 m, rainforest (B.L. Fisher et al.) (CAS); FC Andriantantely, -18.695, 48.81333, 530 m, rainforest (H.J. Ratsirarson) (CAS); FC Didy, -18.19833, 48.57833, 960 m, rainforest (H.J. Ratsirarson) (CAS); FC Sandranantitra, -18.04833, 49.09167, 450 m, rainforest (H.J. Ratsirarson) (CAS); Forêt Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (Malagasy ant team) (CAS); Ile Sainte Marie, Forêt Ampanihy, 14.4 km 52° Ambodifotatra, -16.91117, 49.93917, 10 m, littoral rainforest (B.L. Fisher et al.) (CAS); Mahavelona (Foulpointe), -17.66667, 49.5 in sandy forest (A. Pauly) (CAS); Manakambahiny Atsinanana, -17.75, 48.71667 Primary forest (A. Pauly) (CAS); Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CAS); Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, -15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CAS); Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, -15.17833, 49.635, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CAS); PN Zahamena, Besaky River, -17.75244, 48.85321, 760 m, rainforest (B.L. Fisher et al.) (CAS); PN Zahamena, Onibe River, -17.75908, 48.85468, 780 m, rainforest (B.L. Fisher et al.) (CAS); PN Mananara-Nord, 7.1 km 261° Antanambe, -16.455, 49.7875, 225 m, rainforest (B.L. Fisher et al.) (CAS); Parcelle K9 Tampolo, -17.175, 49.268, 10 m, littoral forest (Malagasy ant team) (CAS); RNI Betampona, Camp Vohitsivalana, 37.1 km 338° -17.88667, 49.2025, 520 m, rainforest (B.L. Fisher et al.) (CAS); RNI Betampona, 34.08 km 332° Toamasina -17.91977, 49.20039, 525 m, rainforest (B.L. Fisher) (CAS); RNI Betampona, 34.1 km 332° Toamasina -17.91614, 49.20185, 550 m, rainforest (B.L. Fisher) (CAS); SF Tampolo, 10 km NNE Fenoarivo Atn. -17.2825, 49.43, 10 m, littoral rainforest (B.L. Fisher) (CAS); Sahafina forest 11.4 km W Brickaville, -18.81445, 48.96205, 140 m, rainforest (B.L. Fisher et al.) (CAS); Tamatave; Andranobolahy; Toamasina Rural, -18, 49, 370 m (CAS). Toliara: 10 km NW Enakara, PN Andohahela, -24.56667, 46.81667, 420 m, rainforest (B.L. Fisher) (PSWC); 13 km NW Enakara, PN Andohahela, -24.55, 46.8, 1250 m, montane rainforest (B.L. Fisher) (CAS); Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina, -24.13993, 47.07418, 540 m, montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); Anosy Region, PN Andohahela, Col de Tanatana, -24.7585, 46.85367, 275 m, rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); Forêt Mandena 8.5 km N Tolagnaro, -24.95267, 47.0025, 20 m, littoral rainforest (B.L. Fisher et al.) (CAS); Isaka-Ivondro; Taolagnaro, -24.8, 46.866667, 41 m (Remy) (CAS); Ivohibe; Mahabo; Betroka, -23.62778, 46.48861, 1552 m (R. Decary) (CAS); Kirindy Forest, plot I, -20.07354, 44.67194, 60 m, dry forest (E. Lokensgard) (CAS); PN Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, -24.76389, 46.75167, 900 m, montane rainforest (Fisher-Griswold Arthropod Team) (CAS); PN Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro, -24.76389, 46.76683, 650 m, rainforest (Fisher-Griswold Arthropod Team) (CAS); Mikea Forest, deciduous dry forest, -22.90367, 43.4755, 30 m, deciduous dry forest (R. Harin’Hala) (CAS); Mikea Forest, spiny forest, -22.91333, 43.48222, 37 m, spiny forest (R. Harin’Hala) (CAS); Tsimelahy-Parcel II, PN Andohahela, transition forest, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 52G). Its colony nests have been found mostly in rotten logs, rotten sticks, and rotting tree stumps, and rarely in dead branches, twigs, and bamboo on the ground. Workers forage frequently on the ground and in leaf litter, rarely on low vegetation.

Figure 52.

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.

Discussion

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 gibber Forel

Figs 32B, 35C, 38A, 53

Camponotus quadrimaculatus var. gibber Forel, 1891: 59. Lectotype minor worker, by present designation, Madagascar, Andrangoloaka (Sikora) AntWeb CASENT0101513 (MHNG) [examined]. Paralectotype. 1 minor worker and 1 alate queen of same data as lectotype but with the following specimen codes: CASENT0101528, CASENT0101537 (MHNG) [examined]. Raised to species by Forel 1891: 215; 1892: 232; Dalla Torre 1893: 232; Emery 1896: 374; Wheeler 1922: 1054; Emery 1925: 122; Bolton 1995: 101. [Combination in Camponotus (Myrmosphincta) Forel, 1914: 273; in Camponotus (Mayria) Emery, 1925: 122].

Additional material examined

Madagascar: Antananarivo: [(de Diversa); Museum Paris, Grandidier 1899]; Mantasoa; Manjakandriana, -19.033333, 47.9166666, 1409 m (CAS); [Madagascar central]; Ambatomanjaka; Miarinarivo, -18.766947, 46.869107, 1343 m (CAS). Antsiranana: Ambondrobe, 41.1 km 175° NW Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest (B.L. Fisher) (CAS); RS Manongarivo 17.3 km 218° SW Antanambao, -14.02167, 48.41833, 1580 m, montane rainforest (B.L. Fisher) (CAS); RS Manongarivo, 14.5 km 220° SW Antanambao, -13.99833, 48.42833, 1175 m, montane rainforest (B.L. Fisher) (CAS); RS Manongarivo, 14.5 km 220° SW Antanambao, -14.00, 48.43167, 1220 m, montane rainforest (B.L. Fisher) (CAS); PN Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, -14.44333, 49.74333, 1325 m, montane rainforest (B.L. Fisher) (CAS); RS Manongarivo, 17.3 km 218° SW Antanambao, -14.02167, 48.41833, 1600 m, montane rainforest (B.L. Fisher) (CAS); PN Marojejy, 11 km NW Manantenina, -14.45, 49.73333, 1875 m, montane rainforest (E.L. Quinter) (CAS). Fianarantsoa: [Hte Sahandrata; Forét prim. de Tsianovoha]; Ambohimitombo; Ambositra, -20.72, 47.45, 1172 m (P.S. Ward) (PSWC): 2 km W Andrambovato, along river Tatamaly, -21.51167, 47.41, 1075 m, montane rainforest (B.L. Fisher et al.) (CAS); 3 km W Ranomafana, nr. Ifandiana, -21.25, 47.41667, 950 m, rainforest (P.S. Ward) (CAS); 43 km S Ambalavao, PN Andringitra, -22.23333, 47, 825 m, rainforest (B.L. Fisher) (CAS); Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CAS); Ambinanindranomena Non Protected Area, 39.16 km SE Ambalavao, -21.96007, 47.29125, 1002 m, Savannah grassland (A. Ravelomanana) (CAS); Ambinanindranomena Non Protected Area, 39.45 km SE Ambalavao, -21.95386, 47.29427, 1069 m, montane rainforest (A. Ravelomanana) (CAS); Ampanenitra Non Protected Area, 41.19 km SE Ambalavao, -21.9652, 47.31001, 1010 m, Savannah grassland (A. Ravelomanana) (CAS); Andrambovato along river Tatamaly, -21.50967, 47.40762, 984 m, cultivated land (tavy) (B.L. Fisher et al.) (CAS); Belle Vue trail, PN Ranomafana, -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin’Hala) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana Belle vue area1.2 km S of PN Ranomafana entrance, -21.2665, 47.42017, 1018 m, rainforest (Rin’Ha, Mike) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana Belle vue area1.2 km S of PN Ranomafana entrance, -21.2665, 47.42017, 1018 m, rainforest (Rin’Ha, Mike) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana, 12 km W of Ranomafana, -21.25083, 47.40717, 1127 m, forest edge, open area (Rin’Ha, Mike) (CAS); JIRAMA water works near river, PN Ranomafana, -21.2485, 47.45217, 690 m, open area near stream (R. Harin’Hala) (CAS); Namorona River at footbridge, PN Ranomafana, -21.25833, 47.42178, 850 m, mixed tropical forest near river, ME Irwin & EI Schlinger (CAS); PN Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, -21.23667, 47.39667, 1150 m, montane rainforest (Fisher, Griswold et al.) (CAS); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); PN Ranomafana, Talatakely, -21.24833, 47.42667, in bamboo forest (CE Griswold, DH Kavanaugh, ND Penny, MJ Raherilalao, JS Ranorianarisoa, J Schweickert) (CAS); RS Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (B.L. Fisher, Sylvain) (CAS); radio tower, PN Ranomafana, -21.25833, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Ranomafana, -21.25, 47.36667 (A. Pauly) (CAS); Ranomafana Nat. Park, Talatakely forest; Sahambavy; Fianarantsoa Rural, -21.4511792, 47.3023894, 1139 m (V.F. Lee, K.J. Ribardo,) (CAS); PN Ranomafana, Talatakely area, 0.4 km WSW of Park Entrance, -21.41667, 47.68333, 900 m, mixed tropical forest (D.H. & K.M. Kavanaugh) (CAS); Vohiparara, -21.23333, 47.36667 (A. Pauly) (CAS); Vohiparara broken bridge, -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin’Hala) (CAS); 23 km E Moramanga, -18.98028, 48.45306, 900 m, tropical dry forest (B.L. Fisher) (CAS). Toamasina: 6.9 km NE Ambanizana, Ambohitsitondroina, -15.56667, 50, 1080 m, montane rainforest (B.L. Fisher) (CAS); 6 km ESE Andasibe (= Perinet), -18.95, 48.46667, 900 m, rainforest (P.S. Ward) (CAS); Bevolota 17.1 km N Andasibe, -18.77071, 48.43164, 995 m, montane rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80424, 48.40081, 968 m, rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.79956, 48.4028, 1058 m, rainforest (B.L. Fisher et al.) (CAS); forêt Didy, Ambatondrazaka, -18.1111503, 48.5107283, 1029 m, rainforest (A. Pauly) (CAS); Mahavelona (Foulpointe), -17.66667, 49.5, Pandanus marsh (A. Pauly) (CAS); Morarano-Chrome, 25 km W forét I, -17.75, 47.98333 (A. Pauly) (CAS); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CAS); PN Andasibe-Mantadia, Forêt de Mantadia, 25.7 km 248° Moramanga, -18.81402, 48.43028, 1040 m, rainforest (F.N. Raharimalala, B. Blaimer) (CAS); Réserve Perinet-Analamazaotra, -18.93333, 48.43333, 950 m, rainforest (D.M. Olson) (CAS). Toliara: 13 km NW Enkara, PN Andohahela, -24.55, 46.8, 1160 m, montane rainforest (B.L. Fisher) (CAS); 13 km NW Enkara, PN Andohahela, -24.56667, 46.81667, 850 m, rainforest (B.L. Fisher) (CAS); Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, -24.13894, 47.06804, 1125 m, rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); Forêt Ivohibe 55.6 km N Tolagnaro, -24.56167, 47.20017, 650 m, rainforest (B.L. Fisher et al.) (CAS); Ifaty 22 km N, -23.18333, 43.61667, 30 m, beach dunes (M.E. Irwin and E.I. Schlinger) (CAS); near road, PN Zombitse, -22.8405, 44.73117, 825 m, spiny deciduous forest (R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 53D). Individual workers forage mostly on the ground and through leaf litter, and rarely on lower vegetation. Nests are typically in rotten logs and rotting tree stumps, but seldom in the ground, in root mats on the ground, in dead twigs above the ground, in moss and leaf litter on live trees, and under tree bark.

Figure 53.

Camponotus gibber A lateral view B head in full-face view C dorsal view of minor worker CASENT0188619 D distribution map.

Discussion

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 gouldi Forel

Figs 31A, 54, 55

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: Dalla Torre 1893: 233; Forel 1897: 201; Forel 1904: 377.

Neotype major worker, by present designation

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 (CAS).

Additional material examined

Madagascar: Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle (Rin’Ha, Mike) (CAS); Antsiranana: Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CAS); SAVA Region, District of Vohemar, Antsahabelela Rain Forest, 9 km SW of Daraina, -13.2505, 49.61667, 182 m, humid Forêt (Mike, Rin’ha) (CAS). Fianarantsoa: 8.0 km NE Ivohibe, -22.42167, 46.89833, 1200 m, montane rainforest (B.L. Fisher, Sylvain) (CAS); PN Ampotoampoto III, 7.91 km NW Ilakaka, -22.62944, 45.189, 919 m, savannah woodland (A. Ravelomanana) (CAS); Anja Reserve, -21.85358, 46.84903, 1085 m, rupicolous vegetation on granite outcrop (B.L. Fisher et al.) (CAS); 1 km E of PN Isalo Interpretive Center, -22.62667, 45.35817, 885 m, dry wash (R. Harin’Hala) (CAS); Forêt d’Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, Uapaca woodland (Fisher, Griswold et al.) (CAS); PN Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CAS); stream area, 900 m E of PN Isalo Interpretive Center, -22.62667, 45.35817, 750 m, open area near stream (R. Harin’Hala) (CAS). Mahajanga: PN Ankarafantsika, Ampijoroa SF, 160 km N Maevatanana, deciduous forest, -16.31944, 46.81333, 43 m, deciduous forest (M. Irwin and Rin’ha Harin’hala) (CAS); Boeny Region, District of Marovoay, PN Ankarafantsika, Ampijoroa SF, 160 km North of Maevatanana on RN 04, -16.31933, 46.81333, 42 m, deciduous forest (Rin’Ha, Mike) (CAS); Forêt Ambohimanga, 26.1 km 314° Mampikony, -15.96267, 47.43817, 250 m, tropical dry forest (B.L. Fisher) (CAS); Forêt de Tsimembo, 8.7 km 336° NNW Soatana, -19.02139, 44.44067, 20 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Melaky Region, District of Besalampy, Marofototra dry forest, 17 km W of Besalampy, -16.72167, 44.42367, 51 m, dry wash in the dry forest (Irwin, Rin’ha) (CAS); Melaky Region, District of Maintirano, Asondrodava dry forest, 15 km N of Maintirano, -17.96533, 44.0355, 6 m, dry forest (Irwin, Rinha) (CAS); PN Ankarafantsika, Ampijoroa SF, 40 km 306° NW Andranofasika, -16.32083, 46.81067, 130 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CAS); Réserve d’Ankoririka, 10.6 km 13° NE Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Station Forestière Ampijoroa, -16.31667, 46.81667, 80 m, tropical dry forest (P.S. Ward) (CAS). Toliara: [Androhomana]; Andranobory; Taolagnaro, -25.18333, 46.63333, 35 m (Sikora); 45 km NE Morondava, -20.05, 44.61667, 30 m, tropical dry forest (P.S. Ward) (CAS); 50 km N Morondava, -20.06667, 44.58333, in primary dry forest (A. Pauly) (CAS); 7.0 km 156° SSE Lavanono, -25.47111, 44.9885, 50 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Androy Region, District of Tsihombe, 74 km S of Tsihombe, RS Cap Ste Marie, -25.58767, 45.163, 36 m, spiny bush (Rin’Ha, Mike) (CAS); Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina, -24.13993, 47.07418, 540 m, montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS);: Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.00667, 46.30333, 85 m, Galery forest (Rin’Ha, Mike) (CAS); Anosy Region, District of Amboasary, PN Andohahela, Parcelle III, Ihazofotsy, 32 km NE Amboasary, -24.83083, 46.53617, 58 m, dry forest, spiny forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.021, 46.3055, 36 m, spiny forest (Mike, Rin’ha) (CAS); Anosy Region, District of Fort-Dauphin, PN Andohahela, Parcelle II, Tsimela, 42 km W of Fort-Dauphin, -24.93683, 46.62667, 176 m, transition forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82505, 46.57811, 90 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82466, 46.60111, 100 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo-Andrefana Region, -23.55275, 43.74471, 45 m, coastal scrub on limestone (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo-Andrefana Region, -23.45314, 43.76448, 20 m, coastal spiny bush on sandy soil (B.L. Fisher, F.A. Esteves et al.) (CAS); Atsimo-Andrefana Region, Antsokay Arboretum, -23.41491, 43.75499, 13 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Fiherenana, -23.17694, 43.96083, 100 m, gallery forest (Frontier Project) (CAS); FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); Forêt de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forét de Kirindy, -20.0671, 44.65723, 50 m (H. Wood & J. Miller) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.06855, 44.65956667, 30 m, tropical dry forest (B.L. Fisher) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.045, 44.66222, 100 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Mahavelo, Isantoria River, -24.75833, 46.15717, 110 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Forêt de Mite, 20.7 km 29° WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Tsinjoriaky, 6.2 km 84° E Tsifota, -22.80222, 43.42067, 70 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Grand Lavasoa, 25.9 km W Tolagnaro, -25.08767, 46.749, 450 m, rainforest edge (B.L. Fisher et al.) (CAS); Mahafaly Plateau, 6.2 km 74° ENE Itampolo, -24.65361, 43.99667, 80 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Makay Mts., -21.26215, 45.17004, 505 m, Barren rock with sparse vegetation, burned grass (B.L. Fisher et al.) (CAS); Makay Mts., -21.34228, 45.18314, 410 m, Gallery forest on sandy soil (B.L. Fisher et al.) (CAS); Makay Mts., -21.20937, 45.25389, 505 m, short scrub, grass, on rocky hill (B.L. Fisher et al.) (CAS); Makay Mts., -21.22284, 45.32477, 490 m, Gallery forest on sandy soil (B.L. Fisher et al.) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza dry forest 07,5 km E of Beroboka, -19.9775, 44.66633, 50 m, dry forest (M. Irwin, Rin’ha) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza galery forest 07 km E of Beroboka, -19.9775, 44.66533, 45 m, Galery forest (M. Irwin, Rin’ha) (CAS); PN Andohahela, Forêt d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, -24.93, 46.6455, 300 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Kirindy Mite, 16.3 km 127° SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, -23.99222, 43.88067, 90 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Mitoho Cave, 6.4 km 77° ENE Efoetse, 17.4 km 170° S Beheloka, -24.04722, 43.75317, 40 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Ranobe, -23.04642, 43.61015, 20 m, spiny forest/thicket (Frontier Wilderness Project) (CAS); Ranobe-plateau, -23.00283, 43.70367, 30 m, grassland (Frontier Wilderness Project) (CAS); RS Beza Mahafaly, Parcel 1, -23.65, 44.63333, 130 m, tropical dry forest (P.S. Ward) (CAS); Réserve Privé Berenty, Forêt d’Anjapolo, 21.4 km 325° NW Amboasary, -24.92972, 46.20967, 65 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Réserve Privé Berenty, Forêt de Bealoka, Mandraré River, 14.6 km 329° NNW Amboasary, -24.95694, 46.2715, 35 m, gallery forest (Fisher-Griswold Arthropod Team) (CAS); RS Cap Sainte Marie, 12.3 km 262° W Marovato, -25.58167, 45.16833, 200 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); 3 km E Itampolo, malaise across path of lower bench of Andrimpano Forest, -24.65783, 43.95617, 45 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km N Ampotaka, malaise on trail in Vitambany gallery forest, -24.65033, 43.96317, 86 m, Gallery forest (M.E. Irwin, Rin’ha) (CAS); 8 km N of Ambohimahavelona village, Ankazomena dry forest, -23.43033, 43.83417, 122 m, dry forest (M.E. Irwin, Rin’ha) (CAS); Ambohimahavelona village 33 km NE of Tulear: Andoharano dry forest, -23.44083, 43.89967, 46 m, dry forest (M.E. Irwin, Rin’ha) (CAS); Ambohimahavelona village 33 km NE of Tulear: private spiny bush, -23.44083, 43.89967, 43 m, spiny bush (M.E. Irwin, Rin’ha) (CAS); PN Andohaela, Tsimelahy, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Andranovato, 5 km SE of Manombo, -22.81806, 43.50217, 18 m, Euphorbia forest (Fisher et al.) (CAS); PN Andohahela, Ihazofotsy - Parcel III,-24.83483, 46.48683, 80 m, tropical dry forest, transition between spiny and dry deciduous forests (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Mikea Forest, -22.90367, 43.4755, 30 m, deciduous dry forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Mikea Forest, -22.91333, 43.48222, 37 m, spiny forest (R. Harin’Hala) (CAS); near ANGAP office, PN Zombitse, -22.8865, 44.69217, 840 m, deciduous spiny forest (R. Harin’Hala) (CAS); near road, PN Zombitse, -22.8405, 44.73117, 825 m, spiny deciduous forest (R. Harin’Hala) (CAS); Parcel I, RS Beza Mahafaly, near research station, -23.6865, 44.591, 165 m, dry deciduous forest (R. Harin’Hala) (CAS); Parcel II, RS Beza Mahafaly, near Bellevue, -23.68983, 44.5755, 180 m, spiny forest (R. Harin’Hala) (CAS); PN Tsimanampetsotsa, Mitoho Forest, malaise on plateau, -24.0485, 43.75233, 150 m, dense dry forest (M.E. Irwin, Rin’ha) (CAS); Tsimelahy - Parcel II, PN Andohahela, transition forest, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Makay, -21.35695, 45.19021, 401 m, xerophylic vegetation (J.M. Bichain) (CAS).

Diagnosis

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.

Description

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.

Figure 54.

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.

Distribution and biology

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. 55D). This species nests mostly in rotten logs, in the ground, and under stones, rarely in rotting tree stumps and rotten sticks. Workers forage most often on the ground and seldom on lower vegetation.

Figure 55.

Camponotus gouldi A lateral view B head in full-face view C dorsal view of minor worker CASENT0121617 D distribution map.

Discussion

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 MHNG, MNHN, NHMB, and MSNG and could not find the type for C. gouldi. Also, the type specimen appears to be absent from the little-known collection of Forel types at Naturmuseum Solothurn (Germann 2017). Morphological characters that define the designated neotype are in accordance with the original description of the former holotype specimen. Since the original type locality was not precisely located within Madagascar, the new locality type has been chosen on the basis of where the species was most often collected across its geographical distribution.

Camponotus hagensii Forel

Figs 33A, 56

Camponotus rubripes r. hagensii Forel, 1886a: 158. Lectotype minor worker, by present designation, Centre de Madagascar (Hildebrandt) AntWeb CASENT0910112 (MHNG) [examined]. Paralectotypes: 2 minor workers and 3 major workers of same data as lectotype but respectively specimen coded as: FOCOL2423, FOCOL2424 (ZMHB), and CASENT0910111 (MHNG), CASENT0104632 and FOCOL2422 (ZMHB), CASENT0101494 (MNHN) [examined]. As subspecies of Camponotus maculatus, Forel, 1891: 27; of Camponotus fumidus, Santschi, 1922: 101. Raised to species by Dalla Torre 1893: 233; Forel 1914: 267; Emery 1920b: 6; Wheeler 1922: 1039; Emery 1925: 92.

Additional material examined

Madagascar: Antananarivo: Central Madagascar; Ambatomanjaka; Miarinarivo, -18.876091, 46.865775, 1343 m (Hildebrandt) (CAS). Antsiranana: 12.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.43333, 1960 m, montane rainforest (B.L. Fisher) (CAS); 12.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.43333, 1985 m, montane rainforest (B.L. Fisher) (CAS); Nosi-Bé du Majunga; Nosy be; Nosibe, -13.315028, 48.25927, 128 m (P. Carié) (CAS); PN Marojejy, 25.4 km 30° NNE Andapa, 10.9 km 311° NW Manantenina, -14.445, 49.735, 2000 m, montane shrubland (B.L. Fisher) (CAS). Fianarantsoa: 36 km S Ambalavao, PN Andringitra, -22.2, 46.96667, 1900 m, montane rainforest (B.L. Fisher) (CAS); 36 km S Ambalavao, PN Andringitra, -22.2, 46.96667, 1975 m, few trees, ericoid thicket (B.L. Fisher) (CAS); 8.5 km SE Antanitotsy, Anjavidilava Forest, -22.16667, 46.96667, 1990 m, Philipia Forest (B.L. Fisher, Sylvain) (CAS); Forest d’Ambalamanakana, -20.73333, 47.2 (A. Pauly) (CAS); PN Andringitra; 8.5 km SE Antanitotsy, -22.16667, 46.96667, 1990 m, montane rainforest (Sylvain) (CAS); Soanierenana IV Non Protected Area, 25.22 km SW Ambositra, -20.72389, 47.10705, 1736 m, savannah grassland (A. Ravelomanana) (CAS); Vohiparara, -21.23333, 47.36667 (A. Pauly) (CAS). Toliara: Anosy Region, Anosyenne Mts, 32.5 km NW Manantenina, -24.14098, 47.03689, 1900 m, montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); near ANGAP office, PN Zombitse, -22.8865, 44.69217, 840 m, deciduous spiny forest (R. Harin’Hala) (CAS); near road, PN Zombitse, -22.8405, 44.73117, 825 m, spiny deciduous forest (R. Harin’Hala) (CAS).

Diagnosis

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.

Description

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.

Distribution and biology

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. 56D). The species has been found nesting in the ground, in root mat layers on the ground, and in rotting tree stumps. Workers have been found foraging frequently on the ground and leaf litter, rarely on low vegetation.

Figure 56.

Camponotus hagensii A lateral view B head in full-face view C dorsal view of minor worker CASENT0191568 D distribution map.

Discussion

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.

Camponotus harenarum sp. nov.

http://zoobank.org/98A3E724-B721-4620-AC59-C6246D6E657D

Figs 5B, 57

Holotype worker

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 (CAS).

Paratype

1 minor worker of same data as holotype but with collection code: BLF09463 and specimen code: CASENT0499217 (CAS).

Additional material examined

Madagascar: Antsiranana: Forêt d’Analabe, 30 km 72° ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher) (CAS).

Diagnosis

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.

Description

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.

Major worker

Unknown.

Distribution and biology

Endemic to Madagascar, C. harenarum is only known from the littoral rainforest of Analabe in Daraina, in the northeastern region of the island (Fig. 57D). This species has been found nesting in rotten logs and foraging on the forest floor and on lower vegetation.

Figure 57.

Camponotus harenarum A lateral view B head in full-face view C dorsal view of holotype minor worker CASENT0499207 D distribution map.

Discussion

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.

Etymology

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 hova Forel

Figs 18A, 58

Camponotus maculatus r. hova Forel, 1891: 35. Syntype workers and queen, Morondava côte ouest de Madagascar (Grandidier) (MHNG); 1 syntype major worker designated as lectotype, by present designation, AntWeb CASENT0101908 (MHNG) [examined]. [First available use of Camponotus rubripes maculatushova Forel, 1886b: 150; unavailable name]. Raised to species by Dalla Torre 1893: 235; Emery 1925: 85; Bolton 1995: 104.

Camponotus hova var. obscuratus Emery, 1925: 85. Syntype workers and male, SW Madagascar (Voeltzkow) (NHMB); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101110 (NHMB) [examined]. Paracletotype major worker with same data as lectotype but specimen coded as: CASENT0101109 (NHMB) [examined]. [First available use of Camponotus maculatus radamae obscurata Forel, 1907: 89; Camponotus maculatus hova obscurior Santschi, 1911a: 132; unavailable names]. Syn. nov.

Additional material examined

Europa Island: Europa Island (Voeltzkow) (MHNG); -22.34775, 40.37041, 10 m, spiny forest on coral (B.L. Fisher) (CAS); -22.33909, 40.38752, 8 m, coastal dune vegetation (B.L. Fisher) (CAS). Juan De Nova Island: -17.04873, 42.71009, 10 m, littoral vegetation (B.L. Fisher) (CAS); -17.06182, 42.72513, 5 m, scrub on coastal karst (B.L. Fisher) (CAS). Madagascar: Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle, (Rin’ha, Fanja) (CAS), Beapombo I Non Protected Area, 22.51 km SW Antsirabe, -20.06892, 47.00404, 1663 m, Savannah grassland (A. Ravelomanana) (CAS). Antsiranana: Antsiranana II Pref: Antsahampano S.-Pref: Montagne d’Ambre. Site MD1, -12.52765, 49.17235, 1049 m, in Commelina regrowth on path next to degraded primary riparian rainforest (D. Lees, R. Ranaivosolo & P. Razafindraibe) (CAS); Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CAS); Nosy Faly, -13.36435, 48.49137, 40 m, open secondary vegetation (B.L. Fisher et al.) (CAS); Andranomatàna, 15.2 km NW Ambilobe, -13.14965, 48.91765, 28 m, sugar cane plantation (B.L. Fisher et al.) (CAS); Forêt d’Antsahabe, 11.4 km 275° W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher) (CAS); Forêt d’Analabe, 30.0 km 72° ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher) (CAS); Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) (CAS); Réserve Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) (CAS); Réserve Analamerana, 28.4 km 99° Anivorano-Nord, -12.74667, 49.49483, 60 m, tropical dry forest (B.L. Fisher) (CAS); RS Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, cultivated land (Fisher, Griswold et al.) (CAS); Sakaramy, -12.44131, 49.22723, 365 m, tropical dry forest (B.L. Fisher et al.) (CAS); Sakaramy, -12.44071, 49.23061, 350 m, grassland (B.L. Fisher et al.) (CAS); Sakaramy, -12.44275, 49.2326, 313 m, tropical dry forest (B.L. Fisher et al.) (CAS); Sakaramy 07 km N of Joffre Ville, -12.33333, 49.25, 360 m, Low rain forest in open area (Mike, Frank, Rin’ha) (CAS); 7 km N Joffreville [camp 2 of Fisher], -12.33333, 49.25, 360 m, in dry forest (R. Harin’Hala) (CAS); PN Montagne d’Ambre [Petit Lac road], -12.533333, 49.166667, 1125 m, rainforest (R. Harin’Hala) (CAS); Montaigne Francais, 150 m, along forested limestone ridge (R. Harin’Hala) (CAS); PN Montagne d’Ambre [1^st campsite], 960 m, rainforest (R. Harin’Hala) (CAS); PN Montagne d’Ambre [lemur trail], 975 m, rainforest (R. Harin’Hala) (CAS); Andilana, Nosy Bé, -13.25, 48.18333, <5 m (D.M. Olson) (CAS). Mahajanga<

﻿Abstract

Keywords

﻿Introduction

﻿Materials and methods

﻿Abbreviation of depositories

﻿Materials

﻿Methods

﻿Measurements

﻿Multivariate statistical analyses

﻿Results and discussion

﻿Multivariate statistical analysis of morphometric data

Taxonomic synopsis of the subgenus Myrmosaga

Morphological diagnosis of the worker caste of Malagasy Myrmosaga

Minor worker

Major worker

Comments

﻿Synoptic list of species of the Malagasy Myrmosaga

﻿Identification key to minor worker caste of the Malagasy Camponotus subgenus Myrmosaga

﻿Species accounts

﻿ Camponotus aina sp. nov.

Holotype worker

Paratype

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

Etymology

﻿ Camponotus aro sp. nov.

Holotype worker

Paratypes

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

Etymology

﻿ Camponotus asara sp. nov.

Holotype worker

Paratypes

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

Etymology

﻿ Camponotus atimo sp. nov.

Holotype worker

Paratype

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

Etymology

﻿ Camponotus aurosus Roger

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

﻿ Camponotus becki Santschi, stat. nov.

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

﻿ Camponotus bemaheva sp. nov.

Holotype worker

Additional material examined

Diagnosis

Description

Distribution and biology

Discussion

Etymology

﻿ Camponotus boivini Forel, stat. rev.

Note

Additional material examined

Diagnosis

Description

Abstract

Introduction

Materials and methods

Abbreviation of depositories

Materials

Methods

Measurements

Multivariate statistical analyses

Results and discussion

Multivariate statistical analysis of morphometric data

Synoptic list of species of the Malagasy Myrmosaga

Identification key to minor worker caste of the Malagasy Camponotus subgenus Myrmosaga

Species accounts

Camponotus aina sp. nov.

Camponotus aro sp. nov.

Camponotus asara sp. nov.

Camponotus atimo sp. nov.

Camponotus aurosus Roger

Camponotus becki Santschi, stat. nov.

Camponotus bemaheva sp. nov.

Camponotus boivini Forel, stat. rev.

Camponotus bozaka sp. nov.

Camponotus cemeryi Özdikmen, stat. nov.

Camponotus cervicalis Roger

Camponotus daraina sp. nov.

Camponotus dufouri Forel

Camponotus gibber Forel

Camponotus gouldi Forel

Camponotus hagensii Forel

Camponotus harenarum sp. nov.

Camponotus hova Forel