[Herpetology • 2026] Nadzikambia nubila, N. evanescens, N. franklinae & N. goodallae • Sky Islands of Mozambique harbour Cryptic Species of Chameleons: Description of Four New Species of Sylvan Chameleons (Squamata: Chamaeleonidae: Nadzikambia Tilbury, Tolley & Branch, 2006)

 A adult male Nadzikambia mlanjensis (Broadley, 1965); B adult male N. baylissi Branch & Tolley, 2010;
C adult male Nadzikambia franklinae sp. nov.; D adult male N. goodallae sp. nov.;
E adult male N. evanescens sp. nov.; F adult female N. nubila sp. nov. 

Tolley & Conradie, 2026

DOI: doi.org/10.3897/vz.76.e178403 

  

Abstract

Several populations of forest-living chameleons in the genus Nadzikambia have been recorded from the montane sky island forests in northern Mozambique. These populations have not been evaluated for their species status, despite the potential for these allopatric populations having diverged at the species level due to vicariance of forest since the mid-Miocene. With only two described species of Nadzikambia, we hypothesised that candidate (new) species occur on each of four additional montane sky islands surveyed. We applied an integrative taxonomic approach to evaluate this, using morphological and genetic data collected from each population. Their distributions were mapped, the morphological dataset was quantitatively analysed using a multivariate analysis, and one nuclear and three mitochondrial genes were sequenced to generate a phylogeny and allele networks. Independent species delimitation analyses were applied to the genetic dataset (mPTP, SpeciesIdentifier, p distances) as supporting evidence for candidate species. By applying integrative taxonomy under the General Lineage Species Concept, we find support for four new species of Nadzikambia. The montane forests where they occur have declined in extent due to slash and burn agriculture and these forest endemics are presumed to be in a proportional decline as their habitat contracts. By examining historical and present-day satellite imagery, we show that all Nadzikambia species have lost significant proportions of their range. Given they do not occur outside these forests, these species are in imminent danger of extinction.

Keywords: Africa, Chamaeleonidae, conservation priority, Critically Endangered, habitat loss, morphological conservatism, reptiles, species declines, species delimitation, taxonomy

 

Life photos of Nadzikambia: A adult male N. mlanjensis (PEM R18445), B adult male N. baylissi (unvouchered specimen),
C adult holotype male N. franklinae sp. nov. (PEM R21165), D adult holotype male N. goodallae sp. nov. (PEM R24394),
E adult holotype male N. evanescens sp. nov. (PEM R24372), F adult paratype female N. nubila sp. nov. (NHMUK 2025.3278).

Nadzikambia franklinae sp. nov.

Namuli sylvan chameleon

Etymology. The new species is named after the British chemist Rosalind Franklin (1920–1958) whose work on X-Ray crystallography, particularly her legendary “photo 51”, revealed the structure of DNA (Franklin and Gosling 1953). Her ground-breaking work subsequently allowed for the field of phylogenetics to develop, decades later. Today, nearly all modern taxonomy is phylogenetically informed, including the description of N. franklinae sp. nov.

Nadzikambia goodallae sp. nov.

Ribáuè sylvan chameleon

Etymology. This species is named after Jane Goodall (1934–2025), an inspirational scientist who lived and worked in Africa throughout her lengthy career. Although her work was dedicated to the study of Pan troglodytes, the Chimpanzee, she spent much of her life living and working in tropical forest, in particular at Gombe National Park, Tanzania. Like her own study species, this chameleon is a forest endemic and the destruction of forest, and other habitats, both at Mount Ribáuè as well as within the home range of P. troglodytes in Central and West Africa is causing forest-living species to decline to the brink of extinction.

Nadzikambia evanescens sp. nov.

Inago sylvan chameleon

Etymology. This species is named Nadzikambia evanescens with the specific epithet from the Latin ‘evanescens’ meaning ‘vanishing’. The name is a present participle that can be used as an adjective or a noun in apposition, and the specific epithet is the same for all genders. The etymology is to highlight the rapidly vanishing forest on Mount Inago and the peril that this species is currently under. The forest has already been reduced to a few small patches, and the uncontrolled conversion of forest to agriculture is continuing. The consequence could be the demise of this endemic forest species, if action is not taken to stop the forest destruction.

Nadzikambia nubila sp. nov.

Chiperone sylvan chameleon

Etymology. This species is named after the “Ciperoni” – the term used locally for the weather that brings heavy clouds and orographic rainfall to the area. The cloud sustains the mid-elevation wet forest on this mountain. The epithet ‘nubila’ is derived from the Latin ‘nubilus’ meaning “cloudy,” and is modified to the feminine form to agree with the feminine gender of the genus Nadzikambia.

 Krystal Tolley and Werner Conradie. 2026. Sky Islands of Mozambique harbour Cryptic Species of Chameleons: Description of Four New Species of Sylvan Chameleons (Squamata: Chamaeleonidae: Nadzikambia Tilbury, Tolley & Branch, 2006). Vertebrate Zoology. 76: 207-246. DOI: doi.org/10.3897/vz.76.e178403 [21 Apr 2026]

[Herpetology • 2024] Brookesia nofy • A New miniaturized Species of Leaf Cchameleon, Genus Brookesia (Squamata: Chamaeleonidae), from a littoral forest fragment in eastern Madagascar

Brookesia nofy

 Rakotoarison, Hasiniaina, Glaw & Vences, 2024

DOI: 10.11646/zootaxa.5506.4.3

 facebook.com/Andolalao

  facebook.com/IUCNchameleons

 

Abstract

The number of species of miniaturized chameleons of the genus Brookesia, subgenus Evoluticauda, known to science has witnessed a dramatic increase over the past 15 years, due to the discovery of multiple microendemic species of very strong genetic divergence. So far, no described Evoluticauda species are known from the littoral forest of Madagascar’s east coast, one of the most threatened habitat types of the island. Here, we report on the discovery of a new species of Evoluticauda occurring in the littoral forest at Ankanin’ny Nofy, a touristic site at about sea level and close to Vohibola forest where probably the same species occurs. The new species, Brookesia nofy sp. nov., is sister to B. ramanantsoai which occurs in mid-altitude and highland forests (800–1300 m a.s.l.) at a similar latitude of eastern Madagascar, but differs from this species by an enormous genetic divergence in both mitochondrial and nuclear genes (8.1–9.4% uncorrected pairwise distance in the 16S rRNA gene, 1.8–2.5% in c-mos), a relatively shorter tail, a different arrangement of cephalic ridges, and probably smaller body size and relatively larger head. The discovery of B. nofy sp. nov. emphasizes the importance of conserving Madagascar’s last remaining fragments of littoral forest.

Reptilia, Squamata, Chamaeleonidae, Brookesia nofy sp. nov., Ankanin’ny Nofy

Andolalao Rakotoarison, Alida Frankline Hasiniaina, Frank Glaw, Miguel Vences. 2024. A New miniaturized Species of Leaf Cchameleon, Genus Brookesia, from a littoral forest fragment in eastern Madagascar.  Zootaxa. 5506(4); 533-547. DOI: doi.org/10.11646/zootaxa.5506.4.3

 facebook.com/Andolalao/posts/10162053722794090

  facebook.com/IUCNchameleons/posts/1056406103159165

[Herpetology • 2024] Rhampholeon bombayi, R. nalubaale, R. msitugrabensis, ... • Taxonomy of the Rhampholeon boulengeri Complex (Sauria: Chamaeleonidae): Five New Species from Central Africa’s Albertine Rift

Rhampholeon plumptrei

 Hughes, Behangana, Lukwago, Menegon, Dehling, Wagner, Tilbury, South, Kusamba & Greenbaum, 2024

DOI: 10.11646/zootaxa.5458.4.1 

 x.com/Daniel_F_Hughes

 

Abstract

In a recent molecular study, the pygmy chameleon Rhampholeon boulengeri Steindachner, 1911 was shown to contain six genetically distinct, but phenotypically cryptic lineages. Phylogenetic analyses of genetic data demonstrated that several well-supported clades occurred in non-overlapping elevational ranges across the Albertine Rift in Central Africa. In order to resolve the taxonomy of the R. boulengeri complex, we examined the morphology of specimens representing all six genetic lineages, including the type specimens. Results supported the notion that the current taxonomy does not reflect species diversity and further uncovered the extent to which morphological differences were dissociated from genetic divergence in this complex. We formally describe five new species of Albertine Rift Rhampholeon, which reflects the species diversity more accurately within the region. All of the species are morphologically conserved and seem to exhibit a pattern of cryptic speciation similar to that observed in the genus and in other chameleon genera. Several of the new species are distributed in adjacent habitats, but occur in parapatry where they are separated by elevation, while species that overlap in elevation are allopatric. At least one of the new species exhibited bone fluorescence from its facial tubercles when examined under ultraviolet light, which is the first published account for the genus. Our results highlight the importance of investigating cryptic diversity using an integrative framework, especially for widespread species that look similar, and the description of these new species reinforces the Albertine Rift as one of the world’s richest biodiversity hotspots.

Reptilia, Biodiversity, Burundi, chameleon, Democratic Republic of the Congo, new species, reptile, Rwanda, Uganda

Rhampholeon bombayi (named or African explorer Sidi Mubarak Bombay)

Rhampholeon plumptrei (named for Andrew Plumptre)

Rhampholeon monteslunae (named for "Mountains of the Moon" aka Rwenzori Mountains)

R. msitugrabensis (named for Albertine Rift )

R. nalubaale (named for Luganda word for "godess")

 in Hughes, Tolley, Behangana, et al. 2018.  

Daniel F. Hughes, Mathias Behangana, Wilber Lukwago, Michele Menegon, J. Maximilian Dehling, Philipp Wagner, Colin R. Tilbury, Trisan South, Chifundera Kusamba and Eli Greenbaum. 2024. Taxonomy of the Rhampholeon boulengeri Complex (Sauria: Chamaeleonidae): Five New Species from Central Africa’s Albertine Rift.  Zootaxa. 5458(4); 451-494. DOI: 10.11646/zootaxa.5458.4.1

  x.com/Daniel_F_Hughes/status/1796219361939632522

 x.com/AndyPlumptre/status/1796570211400990848

Daniel F. Hughes, Krystal A. Tolley, Mathias Behangana, Wilber Lukwago, Michele Menegon, J. Maximilian Dehling, Jan Stipala, Colin R. Tilbury, Arshad M. Khan and Chifundera Kusamba. 2018. Cryptic Diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), A Pygmy Chameleon from the Albertine Rift Biodiversity Hotspot. Molecular Phylogenetics and Evolution. 122; 125-141. DOI: 10.1016/j.ympev.2017.11.015  

[Herpetology • 2022] Rhampholeon colemani, R. rubeho, R. waynelotteri, ... • Cryptic Diversity in Pygmy Chameleons (Chamaeleonidae: Rhampholeon) of the Eastern Arc Mountains of Tanzania, with Description of Six New Species

Rhampholeon colemani,  R. sabini,  

R. rubeho,  R. nicolai, 

R. waynelotteri,  R. princeeai 

Menegon, Lyakurwa, Loader & Tolley, 2022 

 DOI:  10.36253/a_h-12978

  twitter.com/IUCNChameleons 

 Researchgate.net/publication/366657704

Abstract

Previous molecular phylogenetic studies of pygmy chameleons have identified several cases of undescribed cryptic diversity of species, some of which have remained undescribed due to a lack of morphological information. Here, we combine descriptive morphology with principal component analysis, to quantify the overall morphological variation, and phylogenetic analysis to describe six new species of Rhampholeon from the Eastern Arc Mountains, including populations found in the Udzungwa, Rubeho, Nguru, Ukaguru, and Nguu Mountains. From our study we detected only limited morphometric variation between species. We distinguish the new species using genetics, combined with assessment of morphological features, and their geographical distribution. We highlight the threats to pygmy chameleons in East Africa from habitat change and exporting live specimens for the wildlife trade. Based on our understanding, we note a few species that we consider at risk of decline – mainly based on their narrow distribution and their apparent popularity in the export market. This study also further underlines the extraordinary biological value of the relatively small forest patches (less than 3000 km2) of the Eastern Arc, which contain more species of chameleons than any other area in mainland Africa.

Keywords: Afromontane, biodiversity, East Africa, chameleons, new species, reptiles

Rhampholeon uluguruensis Tilbury & Emmrich, 1996

Uluguru Pygmy Chameleon

Rhampholeon moyeri Menegon, Salvidio & Tilbury 2002

Moyer’s Pygmy Chameleon

Rhampholeon beraduccii Mariaux & Tilbury 2006

Beraducci’s Pygmy Chameleon

Rhampholeon acuminatus Mariaux & Tilbury 2006

Nguru Spiny Pygmy Chameleon

Rhampholeon colemani from the Uzungwa Scarp Nature Reserve. 

Rhampholeon sabini from Nguu North Forest Reserve. 

Rhampholeon rubeho from Mafwomero Forest Reserve in the Rubeho Mountains. 

Rhampholeon nicolai from Mamiwa Kisara North Forest Reserve in the Ukaguru Mountains.

Rhampholeon waynelotteri from Kanga Forest Reserve in the Nguru Mountains landscape.

Rhampholeon princeeai from Mkingu Nature Reserve in the Nguru Mountains.

Rhampholeon colemani sp. nov.

Uzungwa Scarp’s Pygmy Chameleon

Etymology: Rhampholeon colemani is named in honour of Carter Coleman, who for more than 25 years has raised funds and campaigned for the conservation of Tanzania’s forests. In 1991 he revived the Tanzania Forest Conservation Group based in Tanzania, and went on to establish the African Rainforest Conservancy in the USA and the Afri-can Rainforest Trust in the United Kingdom, organisations that remain dedicated to conserving Tanzania’s high biodiversity forests

 

Rhampholeon sabini sp. nov. 

Nguu North Pygmy Chameleon

Etymology: The species name is a patronym for Andy Sabin who provides financial support to many organizations and is actively engaged with community and environmental programs around the world. As an extension of his life-long fascination with reptiles and amphibians and dedication to environmental education.

Rhampholeon rubeho sp. nov. 

Rubeho’s Pigmy Chameleon

Etymology: The species is named after the mountain block (Rubeho) where the type series was collected and where it is con-sidered to be restricted. The specific epithet is considered to be a noun in apposition.

Rhampholeon nicolai sp. nov. 

Nicola’s Pigmy Chameleon

Etymology: This species is named after Nicola Colangelo, an entrepreneur and industrialist who worked in Tanzania for most of his life. He supported conservation initiatives and organisations, and before many others argued for the sustainable utilisation of natural resources. He was passionate about conservation for its intrinsic importance but also for a sustainable development of economies, such as low impact tourism activities.

Rhampholeon waynelotteri sp. nov.

Wayne’s Pygmy Chameleon

Etymology: This species is named after and dedicated to Wayne Lotter, in recognition to his ground-breaking work in developing a holistic and strategic intelligence-based approach to antipoaching. He helped successfully reverse the high rates of elephant poaching in Tanzania, during 2010’s. Unfortunately, he died on 16 August 2017 bravely fighting for the cause he was most passionate about. The specific epithet is patronym in the genitive masculine singular.

Rhampholeon princeeai sp. nov. 

Princeeai’s Pygmy Chameleon

Etymology: The species is named after Richard Williams, better known by his stage name Prince Ea, an American activist and inspirational spoken word artist, poet, rapper and filmmaker. From 2014 he has shifted his focus from music to creating motivational and inspirational spoken word films and content covering a wide range of topics such as deforestation and the reckless destruction of our environment for which we are all responsible. The specific epithet is considered to be a noun in apposition.

Michele Menegon, John V. Lyakurwa, Simon P. Loader and Krystal A. Tolley. 2022. Cryptic Diversity in Pygmy Chameleons (Chamaeleonidae: Rhampholeon) of the Eastern Arc Mountains of Tanzania, with Description of Six New Species. Acta Herpetologica. 17(2; 85-113. DOI:  10.36253/a_h-12978

  twitter.com/IUCNchameleons/status/1608487507808735237  

Researchgate.net/publication/366657704_Cryptic_diversity_in_Rhampholeon_of_the_Eastern_Arc_Mountains_of_Tanzania

[Herpetology • 2022] Convergence and Vicariance: Speciation of Chameleons in the Cape Fold Mountains, South Africa, and the Description of Three New Species of Bradypodion Fitzinger, 1843 (Squamata: Chamaeleonidae)

Bradypodion spp.

Tolley, Tilbury & Burger, 2022

DOI: 10.1080/21564574.2021.1998236 

  facebook.com/IUCNChameleons 

ABSTRACT

The mechanisms that underpin ecological speciation, morphological convergence and the evolution of ecological morphotypes (ecomorphs) in squamates have allowed for a better appreciation of the speciation process in chameleons. In particular, attention has been drawn to several populations of chameleons (Sauria, Chamaeleonidae, Bradypodion) from the Cape Fold Mountains, South Africa. Previous work suggested that these populations are genetically divergent, but with strong similarities in phenotype. Using an integrative taxonomic approach that accounts for genetic diversity, habitat and morphology, three of these populations are described as species. One population is from an isolated forest patch and is genetically different at the species level, but morphologically similar to Bradypodion damaranum (Boulenger, 1887) from forested areas in the Knysna region. Although not sister species, the two are in the same clade and probably diverged through vicariance of the forest. Two other populations are from fynbos habitat in adjacent mountain ranges (Tsitsikamma/Langkloof/Kouga mountains and Baviaansklo of Mountains) and are also morphologically similar, but genetically divergent at the species level. These two species are not sister taxa and are not in the same clade yet have a virtually identical phenotype presumably as the result of convergent evolution for the fynbos habitat. Within the context of morphological taxonomy, these populations have been difficult to evaluate. However, when viewed in the context of ecological speciation, convergence and morphological conservatism, the species boundaries are apparent, allowing for them to be described as new taxa.

KEYWORDS: adaptation, Africa, ecomorphology, habitat, new species, reptiles

 

Krystal A. Tolley, Colin R. Tilbury and Marius Burger. 2022. Convergence and Vicariance: Speciation of Chameleons in the Cape Fold Mountains, South Africa, and the Description of Three New Species of Bradypodion Fitzinger, 1843. African Journal of Herpetology.  DOI: 10.1080/21564574.2021.1998236 

  facebook.com/IUCNChameleons/posts/377026687763780

  twitter.com/IUCNchameleons/status/1499596135547219969

A new paper uses an integrative taxonomic approach to assess four chameleon populations from the Cape Fold Mountains of South Africa that are genetically divergent from each other, but with stong phenotypic similarities, describing three new species. Two of those populations (Bradypodion damaranum and Bradypodion venustum sp. nov.) are morphologically similar and from the same clade, and are therefore hypothesized to have diverged through vicariance of the forest. The other two populations (Bradypodion barbatulum sp. nov. and Bradypodion baviaanense sp. nov.) are from different clades but have virtually identical phenotypes presumably as the result of convergent evolution for their fynbos habitat. 

[Herpetology • 2021] Trioceros wolfgangboehmei • A New Chameleon of the Trioceros affinis Species Complex (Squamata, Chamaeleonidae) from Bale Mountains, Ethiopia

Trioceros wolfgangboehmei 

Koppetsch, Nečas & Wipfler, 2021

DOI: 10.3897/zse.97.57297

Photos by Petr Nečas.

Abstract

A new species of chameleon, Trioceros wolfgangboehmei sp. nov., inhabiting the northern slopes of the Bale Mountains in Ethiopia, is described. It differs from its Ethiopian congeners by a combination of the following features: presence of a prominent dorsal crest with a low number of enlarged conical scales reaching along the anterior half of the tail as a prominent tail crest, a casque raised above the dorsal crest, heterogeneous body scalation, long canthus parietalis, rugose head scalation, high number of flank scales at midbody and unique hemipenial morphology. Based on morphological characteristics, phylogenetic discordances of previous studies and biogeographical patterns, this new species is assigned to the Trioceros affinis (Rüppell, 1845) species complex. An updated comprehensive key to the Trioceros found in Ethiopia is provided.

Key Words: Bale Mountains, biogeography, Ethiopia, Great Rift Valley, key, morphology, new species, systematics, taxonomy, Trioceros wolfgangboehmei sp. nov.

Figure 6. Living individual of Trioceros wolfgangboehmei sp. nov. from Dinsho, Ethiopia showing a prominent white temporal spot and dorsolateral longitudinal stripe.

Photo by Petr Nečas. 

Figure 7. Living individual of Trioceros wolfgangboehmei sp. nov. from Goba, Ethiopia, showing a beige ground pattern with slight reddish stripes in the head region around the orbit.

Photo by Petr Nečas. 

Trioceros wolfgangboehmei, sp. nov.

 

Suggested common English name: Wolfgang Böhme’s Ethiopian Chameleon

Diagnosis: 

Trioceros wolfgangboehmei sp. nov. is a small-sized chameleon of the Trioceros affinis species complex (sensu Ceccarelli et al. 2014). It can be distinguished from all other members of the same species complex by the following combination of characters:

 

(1) presence of a prominent and well-developed dorsal crest consisting of a relatively low number of significantly pointed and enlarged conical scales, forming a single row and reaching along the anterior half the tail; 

(2) top of the casque posteriorly raised above the dorsal crest;

(3) heterogeneous body scalation with both small scattered standard scales and enlarged flattened plate-like scales;

(4) long canthus parietalis formed by 9–12 slightly enlarged scales;

(5) rugose head scalation consisting of enlarged scales forming the cranial crests that fill the area between the lateral and temporal crest and the posterior rim of the orbit;

(6) relatively high number of flank scales at midbody (53–59);

(7) relatively short snout-vent length (up to 66 mm);

(8) a unique hemipenial morphology including shallow calyces with smooth margins on the truncus, four pairs of thick, pointed and thorn-like papillae and two pairs of non-serrated rotulae.

Etymology: The specific epithet honours Wolfgang Böhme, senior herpetologist at the Zoological Research Museum Alexander Koenig in Bonn, Germany, for his numerous contributions to research on chameleons, for his outstanding and ongoing herpetological research in general, and, last but not least, for his continuously generous support of the first as well as second author and numerous junior zoologists. The species epithet is a noun in the genitive case.

Figure 10. Living juvenile of Trioceros wolfgangboehmei sp. nov. from Goba, Ethiopia.

Photo by Petr Nečas.

Figure 9. Head of a living Trioceros wolfgangboehmei sp. nov. from Goba, Ethiopia.

Photo by Petr Nečas.

Figure 8. Lateral detail of a living Trioceros wolfgangboehmei sp. nov. from Goba, Ethiopia, showing the heterogeneous body scalation with both small scattered tubercles and enlarged flattened plate-like scales. In this individual the dorsolateral stripe is interrupted and forms a Y-shaped pattern on the flanks.

Photo by Petr Nečas.

Thore Koppetsch, Petr Nečas and Benjamin Wipfler. 2021. A New Chameleon of the Trioceros affinis Species Complex (Squamata, Chamaeleonidae) from Ethiopia. Zoosystematics and Evolution. 97(1): 161-179. DOI: 10.3897/zse.97.57297

[Herpetology • 2018] Cryptic Diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), A Pygmy Chameleon from the Albertine Rift Biodiversity Hotspot

  

in Hughes, Tolley, Behangana, .. et Kusamba, 2018. 

DOI: 10.1016/j.ympev.2017.11.015 

 ResearchGate.net/publication/321474137

Highlights: 

• Species diversity of pygmy chameleons in the Albertine Rift is underestimated.

• Rhampholeon boulengeri is a complex of at least 6 species.

• Diversification via allopatry underlies genetic patterns in genus-level phylogeny.

• The R. boulengeri complex exhibits signatures of parapatric speciation.

• We discuss the importance of investigating cryptic diversity in the montane Afrotropics.

Abstract

Several biogeographic barriers in the Central African highlands have reduced gene flow among populations of many terrestrial species in predictable ways. Yet, a comprehensive understanding of mechanisms underlying species divergence in the Afrotropics can be obscured by unrecognized levels of cryptic diversity, particularly in widespread species. We implemented a multilocus phylogeographic approach to examine diversity within the widely distributed Central African pygmy chameleon, Rhampholeon boulengeri. Gene-tree analyses coupled with a comparative coalescent-based species delimitation framework revealed R. boulengeri as a complex of at least six genetically distinct species. The spatiotemporal speciation patterns for these cryptic species conform to general biogeographic hypotheses supporting vicariance as the main factor behind patterns of divergence in the Albertine Rift, a biodiversity hotspot in Central Africa. However, we found that parapatric species and sister species inhabited adjacent habitats, but were found in largely non-overlapping elevational ranges in the Albertine Rift, suggesting that differentiation in elevation was also an important mode of divergence. The phylogeographic patterns recovered for the genus-level phylogeny provide additional evidence for speciation by isolation in forest refugia, and dating estimates indicated that the Miocene was a significant period for this diversification. Our results highlight the importance of investigating cryptic diversity in widespread species to improve understanding of diversification patterns in environmentally diverse regions such as the montane Afrotropics.

Keywords: Biodiversity, Biogeography, Burundi, Diversification, Democratic Republic of the Congo, Kenya, Molecular systematics, Phylogeography, Rwanda, Uganda

Daniel F. Hughes, Krystal A. Tolley, Mathias Behangana, Wilber Lukwago, Michele Menegon, J. Maximilian Dehling, Jan Stipala, Colin R. Tilbury, Arshad M. Khan and Chifundera Kusamba. 2018. Cryptic Diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), A Pygmy Chameleon from the Albertine Rift Biodiversity Hotspot. Molecular Phylogenetics and Evolution. 122; 125-141. DOI: 10.1016/j.ympev.2017.11.015  

ResearchGate.net/publication/321474137_Cryptic_diversity_in_Rhampholeon_boulengeri_a_pygmy_chameleon_from_the_Albertine_Rift_biodiversity_hotspot

 ssarherps.org/wp-content/uploads/2017/07/Hughes_et_al.pdf

[Herpetology • 2021] Brookesia nana • Extreme Miniaturization of A New Amniote Vertebrate and insights into the Evolution of Genital Size in Chameleons

Brookesia nana 

Glaw, Köhler, Hawlitschek, Ratsoavina, Rakotoarison, Scherz & Vences, 2021 

DOI: 10.1038/s41598-020-80955-1 

  twitter.com/MarkScherz

Abstract

Evolutionary reduction of adult body size (miniaturization) has profound consequences for organismal biology and is an important subject of evolutionary research. Based on two individuals we describe a new, extremely miniaturized chameleon, which may be the world’s smallest reptile species. The male holotype of Brookesia nana sp. nov. has a snout–vent length of 13.5 mm (total length 21.6 mm) and has large, apparently fully developed hemipenes, making it apparently the smallest mature male amniote ever recorded. The female paratype measures 19.2 mm snout–vent length (total length 28.9 mm) and a micro-CT scan revealed developing eggs in the body cavity, likewise indicating sexual maturity. The new chameleon is only known from a degraded montane rainforest in northern Madagascar and might be threatened by extinction. Molecular phylogenetic analyses place it as sister to B. karchei, the largest species in the clade of miniaturized Brookesia species, for which we resurrect Evoluticauda Angel, 1942 as subgenus name. The genetic divergence of B. nana sp. nov. is rather strong (9.9‒14.9% to all other Evoluticauda species in the 16S rRNA gene). A comparative study of genital length in Malagasy chameleons revealed a tendency for the smallest chameleons to have the relatively largest hemipenes, which might be a consequence of a reversed sexual size dimorphism with males substantially smaller than females in the smallest species. The miniaturized males may need larger hemipenes to enable a better mechanical fit with female genitals during copulation. Comprehensive studies of female genitalia are needed to test this hypothesis and to better understand the evolution of genitalia in reptiles.

Brookesia nana sp. nov. in life. 
(A–C) male holotype (ZSM 1660/2012).
(D, E) female paratype (UADBA-R/FGZC 3752).

Molecular phylogeny of specimens in the subgenus Evoluticauda (known as Brookesia minima species group), based on the nuclear CMOS gene (alignment length 847 bp, but only about 400 bp available for all samples) and inferred under the Maximum Likelihood optimality criterion (K2P + G substitution model). Values at nodes are support values from a bootstrap analysis in percent (500 replicates) and are only shown if > 50%. The tree was rooted with B. brygooi (removed for better graphical representation).

Systematics

Order Squamata Oppel, 181115.

Family Chamaeleonidae Rafinesque, 181516.

Subfamily Brookesiinae Angel, 194217.

Genus Brookesia Gray, 186517.

Subgenus Evoluticauda Angel, 1942 (resurrected herein, justification below).

Brookesia nana sp. nov

 

 Diagnosis: A diminutive chameleon species assigned to the genus Brookesia on the basis of its small body size, short tail, presence of rows of dorsolateral tubercles along vertebral column, presence of pelvic spine, and molecular phylogenetic relationships. Brookesia nana sp. nov. is distinguished by the following unique suite of morphological characters: (1) male SVL 13.5 mm, female SVL 19.2 mm; (2) male TL mm 21.6 mm, female TL 28.9 mm; (3) TaL/SVL ratio of 0.51 in male; (4) absence of lateral or dorsal spines on the tail; (5) absence of dorsal pelvic shield in sacral area; (6) presence of distinct pelvic spine; (7) pale brown dorsal colouration with slightly darker markings in life; (8) absence of apical spines on the hemipenis.

Etymology: The specific epithet is the Latin noun nana (meaning female dwarf) in the nominative singular.

Map of northern Madagascar, showing the distribution of species of the subgenus Evoluticauda (known as Brookesia minima group) in this region (only showing records verified by molecular data). Note that B. dentata, B. exarmata, and B. ramanantsoai occur further south and are not included in the map. Orange (dry forest) and green (rainforest) show remaining primary vegetation in 2003–2006.

....

Due to the old divergence and the morphological distinctness of the two clades we here suggest to consider them as different subgenera:

Subgenus Brookesia Gray, 1865 (large-bodied clade)

Type species: Chamaeleo superciliaris Kuhl, 1820.

Contents: Brookesia antakarana, B. bekolosy (attribution tentative), B. betschi, B. bonsi, B. brunoi, B. brygooi, B. decaryi, B. ebenaui, B. griveaudi, B. lambertoni, B. lineata, B. perarmata, B. stumpffi, B. superciliaris, B. therezieni, B. thieli, B. vadoni, B. valerieae.

Distribution: Madagascar.

Subgenus Evoluticauda Angel, 1942 (miniaturized clade, known as B. minima group)

Type species: Brookesia tuberculata Mocquard, 1894.

Contents: Brookesia confidens, B. dentata, B. desperata, B. exarmata, B. karchei, B. micra, B. minima, B. nana, B. peyrierasi, B. ramanantsoai, B. tedi, B. tristis, B. tuberculata.

Distribution: Northern half of Madagascar.

 

Frank Glaw, Jörn Köhler, Oliver Hawlitschek, Fanomezana M. Ratsoavina, Andolalao Rakotoarison, Mark D. Scherz and Miguel Vences. 2021. Extreme Miniaturization of A New Amniote Vertebrate and insights into the Evolution of Genital Size in Chameleons. Scientific Reports. 11, 2522. DOI: 10.1038/s41598-020-80955-1 

  twitter.com/MarkScherz/status/1354734668084637696

[Herpetology • 2020] Rediscovery, Conservation Status and Genetic Relationships of the Malagasy Chameleon Furcifer voeltzkowi (Squamata: Chamaeleonidae)

Furcifer voeltzkowi (Boettger, 1893)

in Glaw, Prötzel, Eckhardt, ... et Vences, 2020. 

 SALAMANDRA. 56(4); 

Abstract

 The chameleon Furcifer voeltzkowi (Boettger, 1893) from northwestern Madagascar was considered to be a synonym of Furcifer rhinoceratus for many decades and was resurrected only recently based on studies of the morphology and osteology of a few male specimens, which were collected more than 100 years ago. However, basic data on this species remain unavailable, including its conservation status, life history, colouration in life, morphology of the female, genital morphology of the male, phylogenetic affinities, and genetic differentiation from F. labordi and other congeners. During a targeted expedition, we rediscovered F. voeltzkowi in its natural habitat close to its type locality, allowing us to fill some gaps of knowledge. Furcifer voeltzkowi is a sexually dimorphic species. The life colouration of males is largely green, whereas that of females is highly variable and can be extremely colourful. Both, morphology and life colouration of males and females show close similarities to F. labordi from west and southwest Madagascar, but also a number of distinct differences (e.g. a larger size of F. voeltzkowi and a smaller rostral appendage in both sexes), enabling a clear distinction of the two species in both sexes. DNA sequences of the nuclear CMOS gene and two mitochondrial markers (16S rRNA and ND4) also confirm that F. voeltzkowi is a distinct species and sister to F. labordi (pairwise differences in the 16S gene 3.5–3.6%). We estimate the distribution range (extent of occurrence) of F. voeltzkowi to comprise ca. 1.000 km2 and suggest that it qualifies as Endangered B1ab(iii) under the IUCN Red List Criteria as its populations are expected to be severely fragmented, and there is continuing decline in the extent and quality of its habitat. Similar to F. labordi, F. voeltzkowi might be an extremely short-living chameleon with a post hatching life span of only several months in the rainy season. The assumed short life might also partly explain why this splendid species got “lost” for many decades, since most roads in its habitat are not accessible in the wet season. 

Key words. Squamata, Chamaeleonidae, morphology, molecular genetics, rediscovery, conservation. 

Figure 2. Males of Furcifer voeltzkowi from Katsepy near Mahajanga in different colour states:
A) slightly stressed; B, C) relaxed (subadult); D, E) displaying.
Note in (A) the injuries on the head probably resulting from intraspecific conflicts. Males differ from those of the sister species F. labordi by larger size, a lower casque, a shorter rostral appendage, darker skin around the scales and a row of white and distinctly enlarged tubercles along the flanks.

Figure 3. Females of Furcifer voeltzkowi from Katsepy near Mahajanga in different colour states:
A) stressed/gravidity colouration; B, C) rather relaxed; D, E) stressed. The females differ from those of the sister species F. labordi by larger size, often by a rudimentary, poorly recognizable rostral appendage, and several details of their colouration (e.g. 2–3 red dorsolateral spots versus only one in F. labordi).

Furcifer voeltzkowi (Boettger, 1893)

Frank Glaw, David Prötzel, Falk Eckhardt, Njaratiana A. Raharinoro, Rojo N. Ravelojaona, Timon Glaw, Kathrin Glaw, Julia Forster and Miguel Vences. 2020. Rediscovery, Conservation Status and Genetic Relationships of the Malagasy Chameleon Furcifer voeltzkowi. SALAMANDRA. 56(4); 342–354.  salamandra-journal.com/index.php/home/contents/2020-vol-56/1996-glaw-f-d-proetzel-f-eckhardt-n-a-raharinoro-r-n-ravelojaona-t-glaw-k-glaw-j-forster-m-vences/file

 globalwildlife.org/press/found-rediscovery-of-lost-chameleon-reveals-reptiles-spectacular-true-colors

 news.mongabay.com/2020/11/a-chameleon-not-seen-in-a-century-reappears-in-a-madagascar-garden