Remaining forests on the Central Highlands of Madagascar-Endemic and endangered aquatic beetle fauna uncovered.

Madagascar is known for its high endemism and as many as 90% of this unique diversity are forest-dwellers. Unfortunately, the forest cover of Madagascar is decreasing at an alarming rate. This decrease can also affect aquatic insects, but our knowledge on aquatic insect diversity and distribution on Madagascar are limited. Although the eastern rainforests are considered the most diverse, the Central Highlands of Madagascar also harbors unique microendemic fauna but has been less studied. Here, we analyze the aquatic Adephaga beetle fauna of three remaining protected forests of the Central Highlands. Diversity, abundance, and uniqueness are compared between and within natural forests and surrounding grasslands. At least 15 undescribed species were found, highlighting the Central Highlands as an important area for endemism. The natural forests and the surrounding grasslands differed significantly in species assemblages. Interestingly, the three remaining forests differed in their assemblages with the geographically more distant Manjakatompo Ankaratra having the most unique fauna but also the highest altitude span. By contrast, the species composition was similar between the peripheral zones of each of the three remaining forests. The similarity of the fauna in the peripheral open habitats illustrates how some local forest endemics are replaced with widespread generalists in degraded habitats. Our study shows that the remaining forests of the Central Highlands of Madagascar are important refuges of unique fauna at high risk of extinction.

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

The CamponotussubgenusMyrmosagasubgen. 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 Myrmopytiasyn. 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.Camponotusmaculatusst.fairmairei Santschi syn. nov., is synonymized under C.boivini. The following are synonymized under C.cervicalis: Camponotuscervicalisgaullei Santschi, syn. nov.; Camponotusperroti Forel, syn. nov.; Camponotusperrotiaeschylus Forel, syn. nov.; Camponotusgerberti Donisthorpe, syn. nov. Camponotusdufouriimerinensis Forel, syn. nov. is a synonym of C.dufouri, Camponotushovavar.obscuratus Emery, syn. nov. is a synonym of C.hova, Camponotusquadrimaculatusopacata Emery, syn. nov. is a synonym of C.immaculatus, Camponotusmaculatusst.legionarium Santschi, syn. nov. is a synonym of C.roeseli, Camponotushovamaculatoides Emery, syn. nov. is a synonym of C.strangulatus. The following are synonymized under C.quadrimaculatus: Camponotuskelleri Forel, syn. nov., Camponotuskellerivar.invalidus Forel, syn. nov., Camponotusquadrimaculatussellaris 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.

Insect morphometry is reproducible under average investigation standards.

Morphometric research is being applied to a growing number and variety of organisms. Discoveries achieved via morphometric approaches are often considered highly transferable, in contrast to the tacit and idiosyncratic interpretation of discrete character states. The reliability of morphometric workflows in insect systematics has never been a subject of focused research, but such studies are sorely needed. In this paper, we assess the reproducibility of morphometric studies of ants where the mode of data collection is a shared routine.We compared datasets generated by eleven independent gaugers, that is, collaborators, who measured 21 continuous morphometric traits on the same pool of individuals according to the same protocol. The gaugers possessed a wide range of morphometric skills, had varying expertise among insect groups, and differed in their facility with measuring equipment. We used intraclass correlation coefficients (ICC) to calculate repeatability and reproducibility values (i.e., intra- and intergauger agreements), and we performed a multivariate permutational multivariate analysis of variance (PERMANOVA) using the Morosita index of dissimilarity with 9,999 iterations.The calculated average measure of intraclass correlation coefficients of different gaugers ranged from R = 0.784 to R = 0.9897 and a significant correlation was found between the repeatability and the morphometric skills of gaugers (p = 0.016). There was no significant association with the magnification of the equipment in the case of these rather small ants. The intergauger agreement, that is the reproducibility, varied between R = 0.872 and R = 0.471 (mean R = 0.690), but all gaugers arrived at the same two-species conclusion. A PERMANOVA test revealed no significant gauger effect on species identity (R 2 = 0.69, p = 0.58).Our findings show that morphometric studies are reproducible when observers follow the standard protocol; hence, morphometric findings are widely transferable and will remain a valuable data source for alpha taxonomy.

Taxonomic revision of the Malagasy Camponotus subgenus Mayria Hymenoptera, Formicidae) using qualitative and quantitative morphology.

The Camponotus subgenus Mayria is revised based on the analysis of both qualitative morphological characters and morphometric traits. The multivariate analysis combined the Nest Centroid (NC)-clustering method and Partitioning Algorithm based on Recursive Thresholding (PART) function 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. In this study, 14 species are recognized, of which seven are newly described: C. lamosy sp. n., C. liandia sp. n., C. mainty sp. n., C. manabo sp. n., C. raina sp. n., C. sada sp. n., C. tanosy sp. n. Four species are redescribed: C. christi Forel, C. dromedarius Forel, C. lubbocki Forel, and C. repens Forel. The following are raised to species and redescribed: C. foersteri Forel stat. n., C. maculiventris Emery stat. n., and C. pulcher stat. n. The following are synonymized under C. foersteri: Camponotus cambouei Forel, syn. n.; C. christi ambustus Forel, syn. n.; C. christi ferrugineus Emery, syn. n. and C. pictipes Forel, syn. n. The following are synonymized under C. lubbocki: Camponotus lubbocki christoides Forel, syn. n. and C. lubbocki rectus Forel, syn. n. Also included in this revision are an illustrated species identification key, taxonomic discussions, images, and distribution maps for each species superimposed with the ecoregions of Madagascar.

Taxonomic revision of the Malagasy Camponotus grandidieri and niveosetosus species groups (Hymenoptera, Formicidae) using qualitative and quantitative morphology.

The Camponotus grandidieri species group and Camponotus niveosetosus species group of the Malagasy region are revised. Species delimitation was inferred from the evidence of both qualitative morphological analysis and multivariate morphometry. The multivariate method combined the Nest Centroid (NC)-clustering method and Partitioning Algorithm based on Recursive Thresholding (PART) function to generate hypotheses about species boundaries (clusters) based on 19 continuous morphological traits of minor workers. The proposed species hypotheses were tested by cumulative cross-validated Linear Discriminant Analysis (LOOCV-LDA) and Principal Component Analysis in a shape space (shape PCA). Morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. Here, eight species are recognized, of which three are newly described and five are redescribed. Four species belong to the Camponotus grandidieri species group: auropubens Forel, efitra n. sp., grandidieri Forel, and maintikibo n. sp.; and four species belong to the Camponotus niveosetosus species group: descarpentriesi Santschi, madagascarensis Forel stat. rev., mita n. sp., and voeltzkowii Forel. Camponotus auropubens aldabrensis Forel and C. olivieri freyeri Santschi are synonymized under C. auropubens. Camponotus grandidieri atrabilis Santschi and C. grandidieri comorensis Santschi are synonymized under C. grandidieri. Illustrated species identification keys for both minor and major castes, taxonomic discussions, images, and distribution maps for each species superimposed on the ecoregions of Madagascar are also provided.

Revision of the Malagasy Camponotus edmondi species group (Hymenoptera, Formicidae, Formicinae): integrating qualitative morphology and multivariate morphometric analysis.

The Malagasy Camponotus edmondi species group is revised based on both qualitative morphological traits and multivariate analysis of continuous morphometric data. To minimize the effect of the scaling properties of diverse traits due to worker caste polymorphism, and to achieve the desired near-linearity of data, morphometric analyses were done only on minor workers. The majority of traits exhibit broken scaling on head size, dividing Camponotus workers into two discrete subcastes, minors and majors. This broken scaling prevents the application of algorithms that uses linear combination of data to the entire dataset, hence only minor workers were analyzed statistically. The elimination of major workers resulted in linearity and the data meet required assumptions. However, morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. Prior species hypotheses and the goodness of clusters were tested on raw data by confirmatory linear discriminant analysis. Due to the small sample size available for some species, a factor known to reduce statistical reliability, hypotheses generated by exploratory analyses were tested with extreme care and species delimitations were inferred via the combined evidence of both qualitative (morphology and biology) and quantitative data. Altogether, fifteen species are recognized, of which 11 are new to science: Camponotus alamainasp. n., Camponotus androysp. n., Camponotus bevohitrasp. n., Camponotus galokosp. n., Camponotus matsilosp. n., Camponotus mifakasp. n., Camponotus orombesp. n., Camponotus tafosp. n., Camponotus tratrasp. n., Camponotus varatrasp. n., and Camponotus zavosp. n. Four species are redescribed: Camponotus echinoploides Forel, Camponotus edmondi André, Camponotus ethicus Forel, and Camponotus robustus Roger. Camponotus edmondi ernesti Forel, syn. n. is synonymized under Camponotus edmondi. This revision also includes an identification key to species for both minor and major castes, information on geographic distribution and biology, taxonomic discussions, and descriptions of intraspecific variation. Traditional taxonomy and multivariate morphometric analysis are independent sources of information which, in combination, allow more precise species delimitation. Moreover, quantitative characters included in identification keys improve accuracy of determination in difficult cases.

Revision of the Malagasy ponerine ants of the genus Leptogenys Roger (Hymenoptera: Formicidae).

Leptogenys is the most diverse ponerine ant genus in the world; it is widespread throughout tropical and subtropical regions and there are over 200 extant species described. Most species have ergatoid queens, and many have falcate, bowed mandibles and are specialists on isopod prey. Here, the Malagasy Leptogenys are revised with 60 species recognized, of which 40 are newly described, 18 redescribed, and two subspecies raised to species rank and redescribed. Included in the revision are a key to species based on the worker caste, geographic distributions, descriptions of intraspecific variation, and notes on natural history. The following species are redescribed: L. acutirostris Santschi, L. alluaudi Emery, L. angusta (Forel), L. antongilensis Emery, L. arcirostris Santschi, L. coerulescens Emery, L. falcigera Roger, L. gracilis Emery, L. grandidieri Forel, L. incisa Forel, L. maxillosa (F. Smith), L. oswaldi Forel, L. pavesii Emery, L. ridens Forel, L. saussurei (Forel), L. stuhlmanni Mayr, L. truncatirostris Forel, and L. voeltzkowi Forel. The following are raised to species and redescribed: L. imerinensis Forel stat. rev., stat. n.; and L. suarensis Emery stat. rev., stat. n. The following are described as new: L. alamando sp. n., L. alatapia sp. n., L. ambo sp. n., L. andritantely sp. n., L. anjara sp. n., L. avaratra sp. n., L. avo sp. n., L. barimaso sp. n., L. bezanozano sp. n., L. borivava sp. n., L. chrislaini sp. n., L. comajojo sp. n., L. diana sp. n., L. edsoni sp. n., L. fasika sp. n., L. fiandry sp. n., L. fotsivava sp. n., L. johary sp. n., L. lavavava sp. n., L. lohahela sp. n., L. lucida sp. n., L. malama sp. n., L. mangabe sp. n., L. manja sp. n., L. manongarivo sp. n., L. mayotte sp. n., L. namana sp. n., L. namoroka sp. n., L. pilaka sp. n., L. rabebe sp. n., L. rabesoni sp. n., L. ralipra sp. n., L. sahamalaza sp. n., L. tatsimo sp. n., L. toeraniva sp. n., L. tsingy sp. n., L. variabilis sp. n., L.vatovavy sp. n., L. vitsy sp. n., and L. zohy sp. n. Most of these species are endemic to the region. Of the endemic species, two are restricted to the Comoros (L. comajojo, L. mayotte), 52 occur only in Madagascar, and two are shared by both islands (L. fiandry, L. gracilis). Three species in the maxillosa group, considered introduced to the region, are recorded from Madagascar, Comoros, Mauritius, Reunion, and Seychelles. Leptogenys stuhlmanni, the only species in the stuhlmanni group, which was collected and first described from Moheli by Forel in 1907, has not been rediscovered and may have gone locally extinct. 

Revision of the Pachycondyla sikorae species-group (Hymenoptera: Formicidae) in Madagascar.

The Malagasy ponerine Pachycondyla sikorae-group is revised and a worker-based key to species is presented. Fourteen species are recognised, of which 13 are described as new. The species group is redefined and divided into two species complexes: the sikorae-complex (P. gorogota sp. n., P. haratsingy sp. n., P. ivolo sp. n., P. maeva sp. n., P. mialy sp. n., P nosy sp. n. and P. sikorae Forel) and the vohitravo-complex (P. agnivo sp. n., P. antsiraka sp. n., P. daraina sp. n., P. rovana sp. n., P. tahary sp. n., P. vohitravo sp. n. and P. zoro sp. n.). All 14 species are endemic to Madagascar and distributed across the rainforests in the east and the transitional humid habitats in the northwest of Madagascar. Distribution maps of each species are included.

Revision of the Pachycondyla wasmannii-group (Hymenoptera: Formicidae) from the Malagasy region.

Defining species limits and describing species of ants are important to identify taxa and habitats with elevated diversity in areas of high conservation priority such as the Malagasy region. The Pachycondyla wasmannii-group is revised in the Malagasy region where eight species are recognized, four of which are new: P. masoala sp. n., P. planicornis sp. n., P. tavaratra sp. n., and P. vazimba sp. n. Four species have been previously described: P. cambouei Forel, P. comorensis (André), P. perroti Forel, and P. wasmannii Forel. Pachycondyla perroti admista Forel is newly synonymized under P. perroti. Pachycondyla cambouei is widespread in eastern Madagascar, morphologically variable, and divided into seven morphotypes. An identification key to species and distribution maps are provided for the genus in the Malagasy region. All species are known only from Madagascar except P. wasmannii, which occurs also on Anjouan of the Comoros Islands.