Malagasy Species Research Articles

Five new species and numerous locality records of Spirobolida millipedes from Madagascar (Diplopoda, Spirobolida, Pachybolidae)

Based mainly on the results of generalized American biodiversity inventory programmes, several samples of Malagasy Spirobolida millipedes became available to study. Based on this material, five new species of Spirobolida are described from Madagascar: four potential microendemic species, Aphistogoniulus amberivery sp. nov. from the Amberivery forest, A. manombo sp. nov., from the lowland rainforest of Manombo, Spiromimus endemicus sp. nov. from the Montagne des Français, and a species from the littoral forest of Tampolo, tentatively placed in the genus Eucarlia Brölemann, 1913, E. tampolo sp. nov.. A fifth species appears to be more widespread and is a morphologically unusual species of the genus Alluviobolus Wesener, 2009, A. omega sp. nov.. Aphistogoniulus manombo sp. nov. was previously described as a population of A. jeekeli Decker & Wesener, 2011. One of the most enigmatic Malagasy millipede species, Spirobolus olympiacus Karsch, 1881, is redescribed based on topotypic material as Colossobolus olympiacus (Karsch, 1881) new combination. Additional locality data is provided for 14 other Spirobolida species, of which 11 are listed on the IUCN Red List: Aphistogoniulus aridus Wesener, 2009, A. diabolicus Wesener, 2009, A. erythrocephalus (Pocock, 1893), A. hova (de Saussure & Zehntner, 1897), Colossobolus semicyclus Wesener, 2009, Spiromimus albipes Wesener & Enghoff, 2009, S. litoralis Wesener & Enghoff, 2009, S. simplex Wesener & Enghoff, 2009, S. triaureus Wesener & Enghoff, 2009, S. univirgatus deSaussure & Zehntner, 1901, Flagellobolus pauliani Wesener, 2009, Dactylobolus bivirgatus (Karsch, 1881), Madabolus maximus Wesener & Enghoff, 2008 and Hylekobolus rufus Wesener, 2009. Fourteen new genetic barcoding COI sequences are provided for nine species: Hylekobolus rufus, Colossobolus semicyclus, Aphistogoniulus amberivery sp. nov., A. aridus, A. diabolicus, A. erythrocephalus, A. hova, Spiromimus univirgatus and S. scapularis Wesener & Enghoff, 2009.

An overview of floral and vegetative evolution in the Asian clade of Bulbophyllum (Orchidaceae)

Background and aims – Bulbophyllum, the largest genus in Orchidaceae, exhibits a diverse morphology in both reproductive and vegetative characters. While trait diversity and evolution has been extensively studied in Malagasy species and within the Cirrhopetalum alliance clade, the evolution of reproductive and vegetative characters at the whole level of the Asian clade remains largely unexplored. Material and methods – We reconstructed the phylogeny of approximately 11% of all Asian Bulbophyllum species using Bayesian inference and maximum likelihood estimation based on nuclear (ITS) and chloroplast (matK, psbA-trnH) DNA sequence data. This phylogenetic framework allowed us to examine the evolution of two vegetative and four floral characters through ancestral state reconstruction. Key results and conclusion – The ancestral character states of the Asian clade of Bulbophyllum include a single leaf, distinct pseudobulbs, multiple-flowered inflorescences, and lateral and dorsal sepals similar in length. One-leaved pseudobulbs evolved into two-leaved pseudobulbs multiple times. Distinct pseudobulbs gave rise to indistinct pseudobulbs twice. Multiple-flowered inflorescences shifted to solitary flowers and 2–3-flowered inflorescences multiple times, with some instances of evolutionary reversal. Lateral sepal elongation also presents a convergent evolutionary scenario.

New species of Dermoergasilus Ho & Do, 1982 (Copepoda: Cyclopoida: Ergasilidae) parasitizing endemic cichlid Paretroplus polyactis (Bleeker) in Madagascar.

Dermoergasilus madagascarensis n. sp. is described from the gills of Paretroplus polyactis, an endemic cichlid fish in Madagascar, using a combined morphological (light microscopy and SEM) and molecular approach (partial 18S rDNA, 28S rDNA, and COI sequences). The new species is characterized mainly by possessing: (i) roughly pentagonal cephalosome; (ii) antennal endopodal segments covered with slightly inflated membrane; (iii) maxillule bearing 2 equally long outer setae and a minute inner seta; (iv) interpodal sternites of swimming legs ornamented with 3–4 rows of spinules; (v) genital segment and first abdominal somite both barrel-shaped; and (vi) a caudal ramus projecting into a digitiform process with inconspicuous terminal seta and bearing 3 terminal setae. The obtained DNA sequences of Malagasy species represent the first molecular data for species of Dermoergasilus. The 28S rDNA phylogeny showed the affiliation of D. madagascarensis n. sp. to Ergasilidae and its sister relationship with cosmopolitan Ergasilus sieboldi von Nordmann, 1832. The first checklist for all species of Dermoergasilus is provided.

Elevation of Three Varieties of the Malagasy Endemic Casearia nigrescens (Salicaceae) to the Species Level

The Malagasy species of Casearia Jacq. (Salicaceae) have no modern taxonomic treatment and present particular difficulties due to their similar flowers. Twentieth-century authors have adopted overly lumping classifications featuring broad circumscriptions of C. nigrescens Tul. Perrier de la Bâthie published three varieties within C. nigrescens sensu Tulasne: varieties onivensis H. Perrier, ovata H. Perrier, and subtrinervia H. Perrier. All of these are from eastern humid forests, farther inland and at higher elevations than is typical of variety nigrescens s. str. Study of the available specimens indicates that all are sufficiently distinct from variety nigrescens that they should be elevated to the rank of species. They are herein recognized as C. onivensis (H. Perrier) Ang. X. Wang & Philpott, C. angustifructa Ang. X. Wang & Appleq., and C. subtrinervia (H. Perrier) Ang. X. Wang & Appleq., respectively. All are rarely collected, and preliminary assessment of their conservation status suggests that all three are Endangered.

Diversity and distribution of Orchidaceae in one of the world's most threatened plant hotspots (Madagascar).

In recent decades, Madagascar has become one of the most important plant hotspots in the world. The country's remaining forests and vegetation are disappearing at an alarming rate, while dozens of new species are discovered each year. Amongst the plant families that have long been studied, Orchidaceae appear as one of the most charismatic, diverse and of high conservation concern. Based on a reviewed, comprehensive herbarium dataset, we have compiled a curated checklist of all orchid species occurring in Madagascar. Based on this complete dataset, we then compiled sampling effort, species diversity distribution and some general statistics on their ecology and IUCN conservation status. We compiled and standardised a global dataset using five public databases as the main data sources, supplemented by the most recent publications. The database contains ~ 10,000 geolocated records collected between 1816 and 2021. We used GIS software and rarefaction methods to examine sampling and diversity patterns. According to our dataset, there are currently 913 orchid species collected in Madagascar, of which 759 orchid species (83.1%) are endemic. Doubling the sampling effort could lead to the discovery of around 100 more species, bringing the total estimated number of orchid species in Madagascar to between 986 and 1048. About one-third (297 species) of all orchid species are known only by type specimens (189 species) or have not been collected in Madagascar for more than 50 years (214 species). Although the raw data show that the Andasibe-Moramanga area would have the highest orchid species concentration, our analysis of the data, adjusted for bias, shows that the centres of orchid diversity in Madagascar are in the Tsaratanàna Strict Nature Reserve and the Ranomafana National Park. Life-form statistics show that 55.0% of orchid species are strict epiphytes. The main flowering period of orchids in Madagascar is between November and March. To date, 84% of the 226 Malagasy orchid species listed in the IUCN Red List are threatened with extinction (CR, EN or VU). Despite geographically uneven coverage, the biodiversity of Malagasy orchids appears to be already well documented. We provide maps corrected for sampling bias that indicate priority areas for future surveys. Upcoming efforts should also focus on rediscovery and conservation of rare and/or threatened species and ensure that the protected area network is well aligned with the distribution of priority species for conservation. Finally, the conservation status of 75% of the orchid species found in Madagascar is not yet known and the inclusion of these species must be a top priority in the coming years.

Taxonomic studies on Malagasy Dalbergia (Fabaceae). V. Eight new large tree species and notes on related Malagasy species

Taxonomic studies on Malagasy Dalbergia (Fabaceae). V. Eight new large tree species and notes on related Malagasy species

Kalanchoe sect. Streptanthae (Crassulaceae subfam. Kalanchooideae), a new two-species section in K. subg. Bryophyllum

A new section, Kalanchoe sect. Streptanthae, with two Malagasy species, K. streptantha and K. bogneri, the only ones included in it, is described in K. subg. Bryophyllum (Crassulaceae subfam. Kalanchooideae).

Phylogeny and infrageneric classification of Clerodendrum (Lamiaceae)

Abstract The infrageneric classification of the genus Clerodendrum (Lamiaceae) has been highly problematic due to different concepts and subdivisions being applied to the treatment of Asian, Australian, and African species. Earlier molecular phylogenetic studies based on Sanger sequencing and limited sampling have indicated that previous morphology-based infrageneric classifications are not congruent with the phylogeny due to morphological convergence in many lineages. Advances in high-throughput DNA sequencing provide more information allowing more robust phylogenetic reconstruction at larger scale. We present the first comprehensive phylogenetic study of Clerodendrum that includes representatives of all previously recognized infrageneric taxa and using targeted sequencing data obtained from the Angiosperms353 enrichment to resolve the phylogenetic relationships. In agreement with previous phylogenetic work, our phylogeny shows that Asian and Australian species form a sister clade to an African Clade. Clerodendrum is monophyletic with the exclusion of some tropical coastal species, including some Australian species, which fall within a clade containing Volkameria and New World genera. We recognize two subgenera: subgen. Clerodendrum for Asian and Australian species and the new subgen. Afroclerodendrum for African and Malagasy species. Our findings support an infrageneric classification of Clerodendrum with a total of 13 sections based on molecular phylogenetic evidence and morphology that clearly accommodate the Asian, remaining Australian, and African species. Of these, we propose three new sections: sect. Albiflora, sect. Fortunata and sect. Megaflora for clades presenting unique morphological characters. The sectional classification and taxonomic consequences are discussed.

A review of Madagascar's derived grasslands: Low palatability following anthropogenic fires may threaten food security

Societal Impact StatementThe 26th United Nations Climate Change Conference emphasised the need to modify practices that negatively impact biodiversity and food security in the context of global change. Following Madagascar's drought‐induced famine, our systematic review supports the theory that grasslands of the Malagasy Central Highland that are subjected to human‐lit fires are anthropogenically derived. Furthermore, these overly frequent fires that characterise much of the Malagasy Central Highland grasslands select poorly palatable grasses. Given the reliance on pastoralism as insurance against crop failure in Madagascar, fire‐dependent practices that degrade rangeland emerge as a threat to food security and biodiversity. Education can mitigate against future humanitarian crises.SummaryFood insecurity is greatest in countries where impacts of global change are predicted to be severe. Many, like Madagascar, rely on livestock‐based pastoralism (and consequently palatable rangelands) for insurance against natural disasters and crop failure. It is recognised that derived grasslands can impact climate and biodiversity. Furthermore, the well‐established palatability‐flammability trade‐off predicts that overly‐frequent fires select increasingly unpalatable, fire‐adapted grassland. The drought‐induced Malagasy famine of 2021 highlights the need to identify factors that threaten food security. Given the ubiquitous practice of rangeland preparation through annual, landscape‐scale human‐lit fires, we evaluate whether Malagasy grasslands are derived and then test for fire‐driven selection of increasingly degraded and unpalatable rangelands across Madagascar's largest grassland system, the Malagasy Central Highland (MCH). We conducted a systematic literature review, evaluating for a palatability‐flammability trade‐off, by determining dominant Malagasy grass species, and then applying functional traits, and palatability ratings to these species. Data were extracted using a suite of relevant search terms, and of 1977 studies identified, 145 were directly relevant to the questions posed. Evidence from the review is compelling for much of the Malagasy highland grassland being derived. Furthermore, Malagasy dominant grass species are fire‐adapted with poor forage‐value, suggesting current burning practices negatively impact both biodiversity and pastoralism. Decreasing rangeland palatability caused by human‐lit fires in a society suffering food insecurity emphasises the need to re‐evaluate pastoralist burning practices. Identifying optimal fire frequencies can avert breaching fire‐induced tipping points to rangeland palatability and the humanitarian crises that may follow.

Malagasy Amphibian Wildlife Trade Revisited: Improving Management Knowledge of the Trade.

Madagascar is a biodiversity hotspot with a long history of trading in its wildlife, especially its hyper-diverse amphibian taxa. Due to globally raised concerns over the conservation of harvested species, CITES was introduced as a global mechanism with which to monitor and regulate the trade. Utilising data collated from the CITES Trade database, this study sought to investigate the trade and CITES' effectiveness in managing the trade with respect to Madagascar. Over a 28-year period, 20 known amphibian species were exported from Madagascar, constituting a total of nearly 271,000 individuals. Formal descriptions of Malagasy amphibian species have increased and continue to increase greatly over time. However, there was no longitudinal relationship regarding the numbers of individuals traded as new species were described. Overall, the number of individuals traded has declined over time, but where assessments were provided by the IUCN Redlist, population declines were reported in all but one species of Malagasy amphibian. Mantella (97.5%) continues to be the predominantly traded genus, with certain, high-conservation-concern, species continuing to be traded. Despite initial concerns over the effectiveness of CITES's actions, after concerted efforts, it appears that CITES' actions were having positive impacts on regulating the trade. However, going forward, concerns remain over the appropriateness of the quotas set and the robustness of their underpinning NDFs. Furthermore, with the increase in the number of recognised species, the potential for incorrect species labelling on the CITES permits increases and requires greater attention.

Three new species of Merma Weise (Coleoptera: Coccinellidae: Epilachnini) from Madagascar.

The herbivorous ladybird genus Merma Weise was regarded as an African endemite, however, recent examination of the type material of the two Malagasy species Epilachna hovana Sicard,1908 [now Merma hovana (Sicard,1908)] and Peralda quadriguttata Sicard, 1909 [now Merma quadriguttata (Sicard, 1909)], revealed they also belong to that genus. Here further three new species of Merma from Madagascar, M. ankarana sp. nov., M. antakotako sp. nov. and M. fisheri sp. nov., are described, diagnosed and illustrated. An updated key to species of Merma from Madagascar is also provided.

A review of small-bodied theropod dinosaurs from the Upper Cretaceous of India, with description of new cranial remains of a noasaurid (Theropoda: Abelisauria)

ABSTRACT Small-bodied theropod dinosaurs are rare on southern landmasses but have been known from India for a century. Excavations by Charles Matley and Durgansankar Bhattacharji in uppermost Cretaceous sediments at Bara Simla, central India in 1917–1919 recovered small theropod vertebral and limb elements originally interpreted as coelurosaurians and separated into at least three species (Compsosuchus solus, Laevisuchus indicus, Jubbulpuria tenuis) based on features that can now be attributed to their serial position in the vertebral column. The comparatively recent discoveries of Noasaurus leali and Masiakasaurus knopfleri from similar-aged rocks in South America and Madagascar, respectively, and advances in basal theropod systematics led to a revised interpretation of most small-bodied Indian theropods as noasaurid abelisauroids. Here we review and redescribe Laevisuchus, Jubbulpuria, and Compsosuchus, including several elements that until now were thought lost, and describe a new partial noasaurid dentary from central India. The dentary bears the characteristic procumbent dentition of Masiakasaurus, which apparently is absent in Noasaurus. Likewise, cervical vertebrae of Laevisuchus more closely resemble those of Masiakasaurus than those of Noasaurus. Despite these similarities, phylogenetic analyses indicate that the balance of character data supports the Indian noasaurid species outside the sister-taxon pairing of South American and Malagasy species. Bones of small-bodied theropods have been recovered exclusively from the youngest Mesozoic localities in India (e.g., Pisdura, Bara Simla); to date they have not been reported from the slightly older localities in western and central India, from southern Indian sites in the Cauvery Basin, nor from the Vitakri Formation of Pakistan.

Kalanchoe ×oberlanderi (Crassulaceae subfam. Kalanchooideae), a horticulturally successful nothospecies with the Malagasy K. blossfeldiana and K. manginii as parents

One of the horticulturally most popular cultivars of Kalanchoe, K. ‘Margrit’s Magic’, (Crassulaceae subfam. Kalanchooideae) in southern Africa was derived from a hybrid between K. blossfeldiana and K. manginii, two Malagasy species. This nothospecies is here described as K. ×oberlanderi. Reference is made to cultivars that likely have been based on K. ×oberlanderi.

A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework

Representatives of Kalanchoe (Crassulaceae subfam. Kalanchooideae) are well-known for having evolved a range of reproductive mechanisms, at least some of which have contributed to certain species and nothospecies having become weeds in places remote from their natural habitats. In addition, kalanchoes have evolved a range of different growth forms in response to a diversity of historical environmental pressures prevalent in the locations where they grow naturally. In terms of growth form, many species are herbaceous, soft- to firm-stemmed perennials that remain small to medium-sized. In some of these same species, variants that are annuals, biennials, or multi-annuals are found. A further group of species are perennial, multi-stemmed shrubs with slightly woody but rather brittle stems and branches. A small group of Malagasy species have woody stems and branches, to the extent that the plants are truly arborescent or nearly so. The distribution of growth form, with special reference to stem and branch morphology, across the four subgenera at present formally recognised in Kalanchoe is reviewed in an evolutionary and classificatory framework, and illustrated.

A DNA barcode reference library for CITES listed Malagasy Dalbergia species.

On Madagascar, the illegal and unsustainable exploitation and illegal international trade of Dalbergia (rosewood) precious woods remain a serious conservation problem. Members of this genus are at high risk of extinction as a consequence of logging, mining, and slash and burn agriculture. Morphological identification of these Malagasy species is difficult in the absence of flowers and fruits, especially in the case of cut trees, sawn wood, and finished product. In this study, we use molecular barcoding to identify the Dalbergia species with the intent to contribute to the control of their illegal trade. Thirty-six Dalbergia samples representing 12 Malagasy species of which 11 have high commercial value, were collected to test the efficacy of a region of the plastid genome (rbcL) and a nuclear-transcribed ITS for barcoding. These widely used markers, as well as DNA barcoding gaps, "best match" and "best close match" approaches, and the neighbor-joining method were employed. All samples were amplified and sequenced using the two markers. Using a single locus, the "best match" and "best close match" approaches revealed that ITS has high discriminatory power within the tested Malagasy species. The combination of rbcL + ITS revealed 100% species discrimination. This study confirms that ITS alone and in combination with chloroplast barcode rbcL allow non-ambiguous identification for the 12 species studied. The results contribute to the development of DNA barcoding as a useful tool to identify Malagasy Dalbergia and suggest that the approach developed should be expanded to all 56 potentially exploited species in reference to international CITES requirements and the sustainable management of valuable resources.

Names validly published in Kalanchoe (Crassulaceae subfam. Kalanchooideae) by Louis-René ‘Edmond’ Tulasne in 1857, with reference to names proposed in sched. by Hilsenberg and Bojer for two Malagasy species, and with notes on the type of the name K. tubiflora

The French botanist Louis-René ‘Edmond’ Tulasne validly published three new names in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in 1857. These are K. eriophylla, K. miniata, and K. floribunda, with the first two of these names being based on material that was collected more than 30 years earlier in Madagascar by Carl Hilsenberg and Wenceslas Bojer, who are jointly credited as having made the first significant plant collections in the interior of the island. The third name, K. floribunda, was published for a species from the Comoros, but is an illegitimate later homonym, for which a lectotype is designated here. The fourth species that Tulasne dealt with in his treatment of Malagasy kalanchoes was included under the name Bryophyllum calycinum, which is now regarded as a synonym of K. pinnata. In the past, Bryophyllum was sometimes accepted as a genus distinct from Kalanchoe, but at present it is treated as K. subg. Bryophyllum. The nomenclature of the three names published in Kalanchoe and the one treated in Bryophyllum by Tulasne is reviewed and discussed in a historical context. The type of the name K. tubiflora, which was collected during a British expedition that lasted from 1822 to 1826 to South and East Africa, is clarified.

How ancient forest fragmentation and riparian connectivity generate high levels of genetic diversity in a microendemic Malagasy tree.

Understanding landscape changes is central to predicting evolutionary trajectories and defining conservation practices. While human-driven deforestation is intense throughout Madagascar, exceptions in areas such as the Loky-Manambato region (north) raise questions regarding the causes and age of forest fragmentation. The Loky-Manambato region also harbours a rich and endemic flora, whose evolutionary origin remains poorly understood. We assessed the genetic diversity of an endangered microendemic Malagasy olive species (Noronhia spinifolia Hong-Wa) to better understand the vegetation dynamics in the Loky-Manambato region and its influence on past evolutionary processes. We characterized 72 individuals sampled across eight forests through nuclear and mitochondrial restriction-associated DNA sequencing data and chloroplast microsatellites. Combined population and landscape genetics analyses indicate that N. spinifolia diversity is largely explained by the current forest cover, highlighting a long-standing habitat mosaic in the region. This sustains a major and long-term role of riparian corridors in maintaining connectivity across these antique mosaic habitats, calling for the study of organismal interactions that promote gene flow.

Season and reproductive activity influence cortisol levels in the Malagasy primate Lepilemur edwardsi.

Throughout the year, wild animals are exposed to a variety of challenges such as changing environmental conditions and reproductive activity. These challenges may affect their stress hormone levels for varying durations and in varying intensities and impacts. Measurements of the glucocorticoid hormone cortisol in the hair of mammals are considered a good biomarker for measuring physiological stress and are increasingly used to evaluate stress hormone levels of wild animals. Here, we examined the influence of season, reproductive activity, sex, as well as body condition on hair cortisol concentrations (HCC) in Lepilemur edwardsi, a small Malagasy primate species. L. edwardsi lives in the seasonal dry forests of western Madagascar, which are characterized by a strongly changing resource availability throughout the year. We hypothesized that these seasonal changes of resource availability and additionally the reproductive cycle of this species would influence HCC of L. edwardsi. Results revealed that hair cortisol concentration of females did not change seasonally or with the reproductive cycle. However, we found a significant increase of hair cortisol levels in males from the early wet season during the early dry season (mating season). This increase is presumably due to changed behavior during the mating season, as sportive lemurs travel more and show aggressive behavior during this time of the year. This behavior is energy-costly and stressful, and presumably leads to elevated HCC. As elevated cortisol levels may impair immune function, L. edwardsi males might also be more susceptible to parasites and diseases, which is unfavorable in particular during a period of low resource availability (dry season).

Transfer of the endemic Malagasy species of Kosteletzkya to Hibiscus and Perrierophytum (Malvaceae)

Hanes, M.M., B. Mashburn & M.W. Callmander (2022). Transfert des espèces endémiques malgaches du genre Kosteletzkya à Hibiscus et Perrierophytum (Malvaceae). Candollea 77: 161–171. En anglais, résumés anglais et français. Madagascar abrite huit espèces qui sont actuellement assignées au genre pantropical Kosteletzkya C. Presl. (Malvaceae). Des études phylogénétiques ont révélé que ces espèces ne sont pas monophylétiques et, au contraire, se répartissent en trois clades distinctes, de rang générique. L'espèce africaine répandue K. adoensis (Hochst. ex A. Rich.) Mast. est la seule espèce du genre Kosteletzkya reconnue à Madagascar, tandis que les sept espèces endémiques malgaches de Kosteletzkya sont transférées à Hibiscus L. et Perrierophytum Hochr. conformément à la morphologie, à la compréhension phylogénétique et à la délimitation générique. Nous proposons donc sept nouvelles combinaisons: Hibiscus retrobracteatus (Hochr.) M.M. Hanes & Callm., Perrierophytum macranthum (Hochr.) M.M. Hanes & Callm., P. madagascariense (Baker) M. Hanes & Callm., P. malvocoeruleum (Hochr.) M. Hanes & Callm., P. reflexiflorum (Hochr.) M.M. Hanes & Callm., P. thouarsianum (Baill.) M.M. Hanes & Callm. et P. velutinum (Garcke) M.M. Hanes & Callm. Quatre nouveaux synonymes sont également proposés et cinq lectotypes sont désignés. Les évaluations du risque d'extinction de ces sept espèces indiquent que deux taxons doivent être classés dans la catégorie «Préoccupation mineure», une «En danger» et quatre «En danger critique d'extinction».

Are Madagascar's obligate grazing-lawns ancient and evolved with endemic herbivores, or recently selected by introduced cattle?

Are Madagascar's obligate grazing-lawns ancient and evolved with endemic herbivores, or recently selected by introduced cattle?