Taxonomy of the Rhampholeon boulengeri Complex (Sauria: Chamaeleonidae): Five New Species from Central Africa’s Albertine Rift

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.

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ISSN 1175-5326 (print edition)
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Accepted by S. Carranza: 6 May 2024; published: 30 May 2024 451
Zootaxa 5458 (4): 451–494
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Copyright © 2024 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5458.4.1
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Taxonomy of the Rhampholeon boulengeri Complex (Sauria: Chamaeleonidae):
Five New Species from Central Africa’s Albertine Rift
DANIEL F. HUGHES1*, MATHIAS BEHANGANA2, WILBER LUKWAGO3, MICHELE MENEGON4,5, J.
MAXIMILIAN DEHLING6, PHILIPP WAGNER7, COLIN R. TILBURY8, TRISAN SOUTH1,11, CHIFUNDERA
KUSAMBA9 & ELI GREENBAUM10
1Department of Biology, Coe College, Cedar Rapids, Iowa 52402, USA.
2Department of Environmental Sciences, Makerere University, P.O. Box 7062, Kampala, Uganda.
�
mbehangana@gmail.com; https://orcid.org/0000-0002-1335-4989
3Department of Environment and Social Safeguards, Uganda National Roads Authority (UNRA), P.O. Box 28487, Kampala, Uganda.
�
Wilber.Lukwago@unra.go.ug; https://orcid.org/0009-0003-3796-0937
4Division of Biology & Conservation Ecology, School of Science & the Environment, Manchester Metropolitan University, Manchester,
UK.
5PAMS Foundation, P.O. Box 16556, Arusha, Tanzania.
�
michele@pamsfoundation.org; https://orcid.org/0000-0002-2366-1771
6Institut für Integrierte Naturwissenschaften, Abteilung Biologie, AG Zoologie, Universität Koblenz-Landau, Universitätsstraße 1,
56070 Koblenz, Germany.
�
dehling@uni-koblenz.de; https://orcid.org/0000-0002-3533-5287
7Allwetterzoo Münster, Sentruper Str 315, D48161 Münster, Germany.
�
wagner@allwetterzoo.de; https://orcid.org/0000-0002-0460-2688
8Department of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland, 7602, South Africa.
�
irgoggaman@gmail.com; https://orcid.org/0000-0002-0329-4513
9Laboratoire d’Herpétologie, Département de Biologie, Centre de Recherche en Sciences Naturelles, Lwiro, République Démocratique
du Congo.
�
chifkusamba@gmail.com; https://orcid.org/0000-0001-7380-7372
10Department of Biological Sciences, University of Texas at El Paso, El Paso, Texas 79968, USA.
�
egreenbaum2@utep.edu; https://orcid.org/0000-0003-1674-1316
11
�
ttsouth21@coe.edu; https://orcid.org/0009-0002-8130-289
*Corresponding author:
�
dhughes@coe.edu; https://orcid.org/0000-0001-5883-5324
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.
Key words: Biodiversity, Burundi, chameleon, Democratic Republic of the Congo, new species, reptile, Rwanda,
Uganda

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Introduction
In the more than 150 years since the first pygmy chameleon, Rieppeleon kerstenii (Peters, 1868), was described from
mainland Africa, a further 27 species have been named—19 in the last 100 years (Uetz et al. 2024)—and there may
be many more additional undescribed species (Tilbury 2018). The pygmy chameleon genus Rhampholeon currently
contains 25 species from sub-montane and montane forests across tropical Africa (Uetz et al. 2024). Rhampholeon
species are forest-floor specialists that exhibit traits closely mimicking a dead leaf, such as a diminutive body size,
brownish body coloration with darker brown streaks that mimic leaf veins, laterally flattened bodies, miniaturized
limb proportions, and a short, functional but non-prehensile tail (Nečas 2004; Nečas & Schmidt 2004; Boistel et al.
2010). The leaf-like phenotype of Rhampholeon is evolutionarily conserved across the genus and a similar appearance
is found among species within the other “pygmy” chameleon genera from both continental Africa (Rieppeleon) and
Madagascar (Palleon and Brookesia). Furthermore, all species in the genus Rhampholeon exhibit considerable
overlap in external characteristics, even between distantly related taxa (Matthee et al. 2004; Branch et al. 2014;
Spawls et al. 2018; Menegon et al. 2022). Phenotypic adaptations to similar environmental selection pressures in
pygmy chameleons seem to underly their highly conserved morphologies, a phenomenon that has concealed many
instances of speciation (Mariaux & Tilbury 2006; Bickford et al. 2007; Tolley & Herrel 2013; Branch et al. 2014;
Menegon et al. 2022). The taxonomic history of Rhampholeon, consequently, has been a complex affair driven by
the difficulty of discovering diagnostic characters among the highly conserved morphologies of dwarfed, terrestrial,
short-tailed, dead-leaf imitators (Nečas & Schmidt 2004). This has been an issue for all four pygmy chameleon
genera, where low interspecific morphological variation and a paucity of indicative physical features have hampered
the characterization and description of species within each genus (Raxworthy & Nussbaum 1995; Glaw et al. 1999;
Glaw et al. 2013; Fisseha et al. 2013; Branch et al. 2014; Menegon et al. 2022). Similar problems have also plagued
taxonomic progress in other chameleon genera, such as Kinyongia (Hughes et al. 2017) and Bradypodion (Tolley
et al. 2022a).
The Austrian zoologist and explorer Rudolf Grauer collected pygmy chameleon specimens over several
expeditions in Central Africa and deposited many of these in the Natural History Museum, Vienna, Austria, under
the care of Franz Steindachner. From Grauer’s specimens, Steindachner (1911) described R. boulengeri, which
he named for the herpetologist George Albert Boulenger from the Natural History Museum, London, United
Kingdom. Of the three type specimens of R. boulengeri (NMW 16000: 1–3), Steindachner recorded from Grauer’s
notes that two of them (both males) came “aus dem Urwald hinter den Sandbergen des nordwestlichen Ufers des
Tanganyikasees” (i.e., from the dense wooded areas [jungle] beyond the sand mountains of the northwest shore of
Lake Tanganyika). The locality of the third type (a female) was less clearly stated, but interpretation of the original
translation indicates that all three most likely came from the same location (W. Denzer, pers. comm.). Grauer also
collected several other pygmy chameleon specimens, including two males and two females from Idjwi Island, South
Kivu, Democratic Republic of the Congo (DRC), and two adult males from a rather vague locality about 90 km west
of the southern shore of Lake Edward at 1600 m elevation. These specimens were reported by Sternfeld (1912),
and apart from slightly longer tails in the latter two males, he concurred that they all represented R. boulengeri.
Steindachner’s (1911) description of R. boulengeri was a short, 174-word account comparing the three specimens
of the new species with Rieppeleon brevicaudatus (Matschie, 1892), a species restricted to the Usambara Mountains
and surrounding area in Tanzania. Steindachner’s description—based on its short tail, relatively smooth dorsal keel,
bicuspid claws, and minor supra-optical horns—would exclude at least 11 other species of Rhampholeon, but cannot
reliably distinguish it from most of the species that were described subsequent to R. boulengeri (Tilbury 2018).
Most pygmy chameleon species are endemic to the small forest fragments from where they were originally
described (Tilbury 2018) and many of these habitats are severely threatened by deforestation (Duveiller et al. 2008;
Di Campo 2017; Aleman et al. 2018; Zeng et al. 2021; Tolley et al. 2022b). Consequently, Rhampholeon is one of the
most threatened groups of chameleons in the world (Tolley et al. 2022b), with 45% (10 out of 22 species evaluated)
listed as Endangered or Critically Endangered (IUCN 2024). Rhampholeon boulengeri, in contrast, is listed as
Least Concern because it is ostensibly distributed across much of tropical Africa from the Congo Basin (DRC)
to Kakamega Forest (Kenya) (Tolley & Plumptre 2014) and can be found across a very wide range of elevation
(Tilbury 2018). Some prior accounts, however, suggested that R. boulengeri is not a single species (e.g., Tolley &
Plumptre 2014; Tilbury 2018), and one study even found three genetic lineages from just three localities, two of
which were < 150 km apart (Tilbury & Tolley 2015). With extensive sampling across the range of R. boulengeri,

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Hughes et al. (2018) confirmed those three lineages plus identified two additional genetically distinct lineages.
Importantly, Hughes et al. (2018) found that the six total lineages (including R. boulengeri [sensu stricto]) did not
overlap with R. hattinghi Tilbury & Tolley, 2015 from the southern Albertine Rift, a species initially considered to
be a conspecific population (Tilbury & Tolley 2015).
Several previous accounts drew attention to R. boulengeri as a candidate species complex in need of taxonomic
revision (e.g., Tilbury & Tolley 2015; Tilbury 2018) and Hughes et al. (2018) further revealed the extent to which
this species is not monotypic. In this paper, we rectify the taxonomic problems present in the R. boulengeri complex
by integrating new morphological data with the molecular data from Hughes et al. (2018) to formally describe five
new species from the Albertine Rift.
Materials and methods
Using the General Lineage Concept (de Queiroz 2007), we diagnose species in the R. boulengeri complex (sensu
Hughes et al. 2018: Rhampholeon sp. 1; Rhampholeon sp. 2; Rhampholeon sp. 3; Rhampholeon sp. 4; Rhampholeon
sp. 5; and Rhampholeon sp. Itombwe [= R. boulengeri]) through the integration of data from multiple sources
(Padial et al. 2010). Specifically, we employ data from morphology, ecology, genetics, and geography to determine
the species diversity within the group. Species comparisons and identifications of pygmy chameleons have been
problematic because of overlapping ranges of numerous characters driven by morphological conservatism in this
group, which is even a challenge among distantly related taxa (Branch et al. 2014; Menegon et al. 2022). Taking an
integrative approach, we describe species-level lineages based on a combination of evidence for populations that
are: (1) allopatric and differentiated genetically/phenotypically; or (2) parapatric and genetically/phenotypically
distinct.
Molecular analyses
To explore genetic variation within the R. boulengeri complex, we refer to net sequence divergences (uncorrected
p-distances) from three gene fragments (16S, ND2, and RAG-1) estimated with MEGA v. 7 (Kumar et al. 2016)
in Table S2 of Hughes et al. (2018) (Supp. Mat. 1). We also refer to phylogenetic results in Hughes et al. (2018),
especially their Figure 4, which is a dated phylogeny based on a concatenated genetic dataset (16S, ND2, and
RAG-1) that consisted of major representatives of Squamata, Sphenodon punctatus, and almost all Chamaeleonidae
including all Rhampholeon species described before 2018 with genetic data on GenBank. This phylogeny was
generated using the program BEAST (Drummond et al. 2012) with five concurrent runs of 100 million generations
each, trees sampled every 5000 generations, and 10% discarded as burn-in. See Hughes et al. (2018) for more
details.
Morphological analyses
Specimens examined for this study were collected from multiple localities throughout the Albertine Rift in Central
Africa, and at least one locality in East Africa. We follow the museum acronyms of Sabaj (2020, 2022) for specimen
numbers. Morphometric data were recorded from preserved specimens with vernier calipers with the aid of a Zeiss
Stemi 2000-C stereomicroscope. Color descriptions are based on color photographs in life, personal observations,
and field notes. Sex was determined by internal examination of gonads, everted hemipenes, or the presence of
hemipenal bulges distal to the vent.
Measurements (± 0.01 mm) were taken from the right side of the body. To reduce potential errors derived from
inter-observer differences (Yezerinac et al. 1992), all specimens were measured by DFH, except the syntypes for
R. boulengeri and R. affinis, which were measured by PW. Morphometric and meristic data, and their associated
abbreviations were modified from Branch & Tolley (2010) and Greenbaum et al. (2012): snout–vent length (SVL)
from tip of snout to anterior edge of vent; tail length (TL) from tip of tail to posterior edge of vent; total length
(ToL) from tip of snout to tip of tail; head length (HL) from superior tip of casque to tip of snout; head width (HW)

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measured at widest point just posterior to eyes; head height (HH) from rictus (i.e., commissure) of jaw to superior
tip of casque; mouth length (ML) from tip of rostral to rictus; casque–eye length (CE) measured diagonally from
posterior margin of orbit to superior tip of casque; snout length (SL) from tip of snout to anterior margin of orbit; eye
diameter (ED) measured horizontally at center of eye; cranial crest gap (CC) measured across the crown between
raised supraorbital crests at mid-eye; inter-limb length (IL) from axillary to inguinal attachments of limbs; forelimb
length (FLL) from elbow to wrist; hind limb length (HLL) from knee to heel; rostral process (RP) length from base
to tip; and two meristic characters, including upper labial tubercles (UL) to posterior margin of orbit; and lower
labial tubercles (LL) to posterior margin of orbit. We note that some specimens were collected for different purposes
(e.g., Hughes et al. 2016), thus not all traits were available to be measured from all specimens, which inevitably
influenced sample sizes across analyses.
The following statistical analyses were conducted in R (R Core Team 2021) using the RStudio interface
(RStudio Team 2021). Mean, standard deviation, and range of mensural and meristic characters were calculated
for each group indicated by the phylogeny. We note that using raw measurements can bias downstream analyses
that compare datasets among lineages because ontogenetic size variation and sexual dimorphism often exist among
measured specimens, and thus removing the effects of size prior to morphometric analyses is crucial. Recently,
Chan & Grismer (2022) showed that the assumptions of isometry and homogeneity of slopes were violated with
ratios and residuals for size-correcting data, thus rendering inappropriate these two standard methods to correct for
size differences. Consequently, we elected to use Thorpe’s (1975) allometric growth model to remove the effects of
body size (Reist 1985) using the package GroupStruct (Chan & Grismer 2022) as it was shown to not violate the
underlying statistical assumptions commonly plaguing the other approaches mentioned above (Chan & Grismer
2022). We relied on the phylogenetic results of Hughes et al. (2018) as a priori evidence for using lineage-specific
body size means for size-correcting measurements. To explore morphological differences between the candidate
species, we followed the protocol outlined by Chan & Grismer (2021) for statistical comparisons among new
species. Specifically, we first tested for sexual dimorphism, checked the size-corrected data for normality, then
employed either parametric (ANOVA) or non-parametric (Kruskal-Wallis) tests of mean differences using variance,
and finally, Tukey’s post hoc comparisons if differences were detected among the lineages. To further examine
whether Rhampholeon from these populations exhibit morphological differences, we used the size-corrected values
as input for a Principal Component Analysis (PCA) and generated 95% confidence ellipses to visually inspect for
overlap among lineages, which we analyzed using both the full morphological dataset and separately by sex. All
variables had communalities > 0.5, a varimax rotation was used, and PCs with eigenvalues > 1.0 were extracted. We
recognized statistical significance at P ≤ 0.05.
Results
Geography and elevation
Genetically distinct lineages exist either allopatrically, or parapatrically where they are separated by elevation
(Figs. 1–3). For example, R. sp. 5 is allopatric with respect to all other lineages as its distribution does not overlap
geographically at either of the sites where it is currently found (Fig. 2), whereas R. sp. 2, R. sp. 3, and R. boulengeri
exist parapatrically at non-overlapping elevations in the Itombwe Natural Reserve, DRC (Fig. 3).
Sequence divergence
Uncorrected p-distances between the six genetic lineages were generally high, while intraspecific values were
consistently lower (Supp. Mat. 1). Using the most variable locus (ND2), p-distance values between lineages and R.
boulengeri ranged from 4.2–5.9% for R. sp. 1; 5.9–6.2% for R. sp. 2; 3.2–4.4% for R. sp. 3; 5.1–5.2% for R. sp. 4;
and 5.1–6.4% for R. sp. 5.

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Sexual dimorphism
Preliminary analyses revealed that males and females were strongly dimorphic in important linear traits, including
SVL (t = 4.42, df = 129, P < 0.001) and TL (t = -5.03, df = 129, P < 0.001). Furthermore, boxplots of morphological
traits separated by sex further showed the extent to which the six lineages exhibited sexual-size dimorphism (Fig.
4). As a result, we chose to analyze the sexes separately in the following statistical comparisons among the lineages
using size-corrected data.
Morphological comparisons—Females
The following morphological traits were normally distributed: HL (Shapiro–Wilk statistic [W] = 0.97, P = 0.09),
HH (W = 0.98, P = 0.31), ML (W = 0.98, P = 0.15), CE (W = 0.99, P = 0.99), ED (W = 0.99, P = 0.99), FLL (W =
0.98, P = 0.33), and HLL (W = 0.97, P = 0.05). The following morphological traits were not normally distributed:
SVL (W = 0.92, P < 0.001), TL (W = 0.96, P = 0.02), HW (W = 0.95, P = 0.004), SL (W = 0.95, P = 0.004), CC (W
= 0.96, P = 0.02), and IL (W = 0.95, P = 0.003).
Overall, the six lineages exhibited significant differences in SVL (χ2 = 15.4, df = 5, P = 0.009), HL (F5,13 = 16.9,
P < 0.001), ML (F5,13 = 6.1, P = 0.003), SL (χ2 = 37.5, df = 5, P < 0.001), ED (F5,13 = 8.7, P < 0.001), HLL (F5,13 = 3.9,
P = 0.02), and IL (χ2 = 47.3, df = 5, P < 0.001).
Post-hoc comparisons identified differences in SVL between R. sp. 5 and R. sp. 3 (P = 0.044), R. sp. 5 and R.
sp. 2 (P = 0.002); HL between R. sp. 2 and R. sp. 1 (P < 0.001), R. sp. 2 and R. sp. 3 (P < 0.0001), R. sp. 2 and R. sp.
4 (P = 0.002), R. sp. 2 and R. sp. 5 (P < 0.0001); ML between R. sp. 2 and R. sp. 1 (P = 0.022), R. sp. 2 and R. sp. 3
(P = 0.039), R. sp. 2 and R. sp. 5 (P < 0.001); SL between R. sp. 5 and R. sp. 1 (P = 0.014), R. sp. 2 and R. sp. 3 (P =
0.006), R. sp. 2 and R. sp. 5 (P < 0.001), R. sp. 5 and R. sp. 3 (P = 0.04); ED between R. sp. 3 and R. sp. 2 (P = 0.009),
R. sp. 5 and R. sp. 1 (P = 0.006), R. sp. 5 and R. sp. 2 (P < 0.001), R. sp. 5 and R. sp. 3 (P = 0.037); HLL between R.
sp. 3 and R. sp. 2 (P = 0.028), R. sp. 5 and R. sp. 2 (P < 0.001); and IL between R. sp. 2 and R. sp. 1 (P = 0.016), R.
sp. 5 and R. sp. 1 (P < 0.0001), R. sp. 3 and R. sp. 2 (P < 0.001), R. sp. 4 and R. sp. 2 (P = 0.007), R. sp. 5 and R. sp.
2 (P < 0.0001), R. boulengeri and R. sp. 2 (P < 0.001), R. sp. 5 and R. sp. 3 (P < 0.001), and R. sp. 5 and R. sp. 4 (P
< 0.001). Lastly, at least one comparison approached significance: SL between R. sp. 2 and R. sp. 1 (P = 0.054).
Morphological comparisons—Males
The following morphological traits were normally distributed: TL (W = 0.96, P = 0.11), HW (W = 0.97, P = 0.24),
ML (W = 0.99, P = 0.97), CE (W = 0.97, P = 0.31), SL (W = 0.97, P = 0.16), ED (W = 0.96, P = 0.065), CC (W =
0.97, P = 0.18), FLL (W = 0.96, P = 0.12), HLL (W = 0.98, P = 0.44), and IL (W = 0.98, P = 0.67). The following
morphological traits were not normally distributed: SVL (W = 0.95, P = 0.048), HL (W = 0.94, P = 0.018), HH (W
= 0.87, P < 0.001), and CC (W = 0.96, P = 0.016).
Overall, the six lineages differed in SVL (χ2 = 9.5, df = 4, P = 0.05), CC (χ2 = 9.3, df = 4, P = 0.05), SL (F4,11 =
3.4, P = 0.05), and IL (F4,11 = 9.7, P = 0.001).
Post-hoc comparisons identified differences in TL between R. boulengeri and R. sp. 1 (P = 0.012), R. boulengeri
and R. sp. 3 (P = 0.018); SL between R. sp. 5 and R. sp. 1 (P = 0.003), R. boulengeri and R. sp. 5 (P = 0.001); and IL
between R. sp. 5 and R. sp. 1 (P < 0.001), R. sp. 5 and R. sp. 3 (P < 0.001), R. sp. 5 and R. sp. 4 (P = 0.031), and R.
boulengeri and R. sp. 5 (P < 0.001). Lastly, several comparisons approached significance, including HH between R.
boulengeri and R. sp. 4 (P = 0.062) and CC between R. boulengeri and R. sp. 1 (P = 0.063).
Principal Component Analysis
Analysis of 13 continuous characters extracted two principal components that accounted for 50.3% of the variation
in the full data set, 51.4% in the female-only data, and 49.6% in the male-only data (Figs. 5–6; Table 1). In the full
data set, the first component (PC1) loaded highest for HL and HLL, and PC2 for TL and IL. In the female-only
analysis, PC1 loaded highest for HL and HLL, and PC2 for CC and IL. In the male-only analysis, PC1 loaded highest

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FIGURE 1. Bayesian phylogeny of the Rhampholeon boulengeri complex from the Albertine Rift, Central Africa, adapted from
Figure 4 of Hughes et al. (2018). Posterior probabilities ≥ 95% are denoted by filled circles adjacent to nodes.

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FIGURE 2. Map of the Albertine Rift, Central Africa, showing sampling localities for specimens of the Rhampholeon boulengeri
complex examined in this study. For clarity, we show color-coded circles that indicate the number of specimens examined from
that general locality. Underlined numbers indicate that a specimen from that area also included genetic data analyzed in Hughes
et al. (2018). Two specimens (UTEP 21704–21705) from DRC not shown (see text for details). For a detailed map of localities
representing R. boulengeri (sensu lato) see Tilbury (2018). The color scheme corresponds to clades of the species complex and
is retained throughout all figures: Green = Rhampholeon sp. Itombwe (R. boulengeri); pink = Rhampholeon sp. 1 (R. plumptrei
sp. nov.); blue = Rhampholeon sp. 2 (R. nalubaale sp. nov.); yellow = Rhampholeon sp. 3 (R. bombayi sp. nov.); purple =
Rhampholeon sp. 4 (R. msitugrabensis sp. nov.); red = Rhampholeon sp. 5 (R. monteslunae sp. nov.).

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FIGURE 3. Elevational zonation of the Rhampholeon boulengeri complex from the Albertine Rift, Central Africa. Each color-
coded circle corresponds to an individual specimen. The top panel shows the elevational distribution by species and the lower
panel is grouped by specific localities. The number enclosed in the larger circle indicates the number of specimens from the
same elevation.

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FIGURE 4. Box plots showing variation in external traits of the Rhampholeon boulengeri complex from the Albertine Rift,
Central Africa. Sample sizes for each species are denoted in the bottom right quarter. Males and females are separated to show
degree of sexual-size dimorphism across members of the complex. All measurements are in mm. Plots show the median,
1st and 3rd quartiles, minimum and maximum inter-quartile ranges, and outliers. Refer to text for description of the linear
measurements.

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FIGURE 5. Bivariate plot of the first two principal components from an analysis of 13 size-corrected measurements using all
specimens for the Rhampholeon boulengeri complex from the Albertine Rift, Central Africa. Loadings and percent variance
explained are presented in Table 1. Lectotype and paralectotypes of R. boulengeri are indicated by green stars. Numbers indicate
specific specimens used in the analysis.

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FIGURE 6. Bivariate plots of the first two principal components from an analysis of 13 size-corrected measurements from the
Rhampholeon boulengeri complex using female (A) and male (B) specimens from the Albertine Rift, Central Africa. Loadings
and percent variance explained are presented in Table 1. Lectotype and paralectotypes of R. boulengeri are indicated by green
stars.
TABLE 1. Principal components analysis (PCA) comparing lineages of the Rhampholeon boulengeri complex with
allometry-corrected morphometric data. Eigenvalues, percent variance, and loadings are shown for the first two principal
components for all specimens, females only, and males only. See Materials and Methods for abbreviations.
All specimens (Fig. 5) Females only (Fig. 6A) Males only (Fig. 6B)
PC1 PC2 PC1 PC2 PC1 PC2
SVL -0.2229758 0.43147509 0.2714242 0.46983758 -0.11068331 -0.46987481
TL -0.4330686 -0.55176432 0.3220499 -0.50900386 -0.65387383 0.41583502
HL -0.8869016 0.12008421 0.9111892 0.09167883 -0.84072803 -0.13081205
HW -0.6942403 -0.10063198 0.6698612 -0.19256751 -0.72299616 -0.22532276
HH -0.4443885 -0.07160585 0.5128073 -0.24785180 -0.32586548 -0.44946950
ML -0.6747938 0.23444900 0.7151186 0.18753035 -0.60065762 -0.01475701
CE -0.5985663 -0.34383425 0.5990225 -0.26903600 -0.62653162 0.21342405
SL -0.6202347 0.34639378 0.6738071 0.28662747 -0.50342321 -0.55558941
ED -0.6795514 0.20253073 0.7440205 0.16115660 -0.61132961 0.02222016
CC -0.4770076 -0.52616822 0.4033642 -0.53023346 -0.64591583 0.03336890
IL -0.3515286 0.73225873 0.4776726 0.64216244 -0.09462887 -0.79414753
FLL -0.7065760 -0.13113648 0.7267426 -0.14663966 -0.66904022 0.31617600
HLL -0.7951229 -0.10151180 0.8062432 -0.07854193 -0.78561259 0.23265436
Eigenvalue 4.85 1.70 5.17 1.52 4.65 1.80
Proportion 37.3% 13.0% 39.7% 11.7% 35.7% 13.9%

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for HL and HLL, and PC2 for IL and SL. Plots of the first two PCs indicated lineage-specific groupings across all
three data sets (Figs. 5–6), several of which demonstrated non-overlapping 95% ellipses among the lineages. In the
analysis using the full data set, R. sp. 2 and R. sp. 5 did not overlap in 95% ellipses with any other species, and R.
sp. 1 and R. sp. 3 did not overlap with R. boulengeri. In fact, only R. sp. 4 shared 95% ellipses with R. boulengeri
in the full PCA. In the sex-specific PCAs, females showed separation in 95% ellipses for species R. sp. 5 and R. sp.
2, and males showed separation for R. sp. 5 and R. sp. 1. The 95% ellipses of the remaining lineages overlapped
in these sex-specific analyses to the extent that would be expected in a group with such extreme morphological
conservatism.
Taxonomy
The members of this complex are phenotypically conserved, and thus morphological traits reported in historical
accounts insufficiently delimited species boundaries in this group. The longstanding recognition of R. boulengeri
as a single species can be explained by limited sampling across its geographic range, lack of sufficient diagnostic
characters, and absence of integrative approaches to delimit species boundaries. Although Steindachner’s (1911)
description of R. boulengeri was published just three years after Grauer collected the specimens in 1908, the type
locality was designated in rather vague terms. Tilbury & Tolley (2015) considered the Itombwe Plateau as the type
locality because Grauer was known to have collected specimens in 1908 from the forests of the Itombwe Plateau,
which is located to the northwest of Lake Tanganyika. Other authors (e.g., Tilbury 2018) have also suggested
this location as the type locality, and genetic data presented by Hughes et al. (2018) furthered the notion that the
species R. boulengeri Steindachner, 1911 may be endemic to the plateau (“R. sp. Itombwe” = R. boulengeri [Hughes
et al. 2018]). Based on the molecular findings of Hughes et al. (2018), morphological results presented above,
and qualitative differences explained below, we describe five new species that were historically considered to be
populations of R. boulengeri.
Systematics of the Rhampholeon boulengeri complex
Family Chamaeleonidae Gray, 1825
Genus Rhampholeon Günther, 1874
Rhampholeon boulengeri Steindachner, 1911
Boulenger’s pygmy chameleon
Chresonymy.
Rhampholeon spectrum boulengeri—Loveridge 1951 (fide Wild 1994)
Rhampholeon boulengeri—Fischer 1996, Nečas 2004 (partim), Matthee et al. 2004 (partim), Tilbury 2010 (partim), Glaw 2015
(partim), Tilbury 2018 (partim), Menegon et al. 2022 (partim)
Rhampholeon sp. Itombwe—Hughes et al. 2018
Original diagnosis (verbatim, translated from German by JMD). In body shape matching Rh. brevicaudatus
(Matschie) [now Rieppeleon brevicaudatus (Matschie, 1892)], but with a short, small, conic rostral process, an
anteriorly concave interorbital ridge and without skin fold from the eye to the tail tip. Tail length about one-fifth of
the total length. Apart from the two-pointed claw no spine on fingers and toes. Eyelid edge more or less distinctly
triangularly acuminate. Temporal ridge and parietal crest well developed. Interorbital edge moderately concave,
the two low eyelid processes connected through a transversal ridge. Helmet low, the rear end merging without
disruption into the back. Rump high like in Rh. brevicaudatus. Dorsal crest weakly toothed. Scalation non-uniform,
the slightly larger tubercles, in particular on the extremities, spine-like pointed.
Head, chest and outer face of extremities, occasionally also the larger or smaller front half of the rump deep
black-brown, the remaining part of the rump brownish white. The underside of the tail, which is thickened in its

FIVE NEW RHAMPHOLEON FROM CENTRAL AFRICA Zootaxa 5458 (4) © 2024 Magnolia Press · 463
basal part, especially in males, is in cross-section flat in males. Three specimens, among them two males, from the
natural forest beyond the sand mountains of the northwestern shore of Lake Tanganyika.
Total length of the largest male 65 mm, tail length somewhat more than 12 mm, greatest rump height 23 mm,
head length 13 mm, greatest head width in the eye region 7.5 mm. Only the outermost tail tip is prehensile.
Original syntypes. Two adult males (NMW 16000-1, NMW 16000-2) and one adult female (NMW 16000-3),
collected by Rudolph Grauer in 1908. Type locality: “aus dem Urwald hinter den Sandbergen des nordwestlichen
Ufers des Tanganyikasees” (i.e., from the dense wooded areas [jungle] beyond the sand mountains of the northwest
shore of Lake Tanganyika), further defined as “forest on the Itombwe Plateau” in eastern DRC by Tilbury & Tolley
(2015), and subsequently upheld by others (e.g., Hughes et al. 2018; Tilbury 2018). According to JMD, the term
“Sandberge” is a misreading of [Graben-] “Randberge” (“rift-flank mountains”), a term used to describe where
many other specimens were collected by Rudolf Grauer in the same area around the same time. The capital letters
“S” and “R” are similar to each other in the style of German handwriting used at that time known as Kurrentschrift.
All other specimens collected by Grauer have the locality as “Randberge.” One of the few localities in that area
specified by Grauer is “Ugoma,” which is where he collected one of the syntypes (NMW 8020) of Chamaeleon
graueri (= Trioceros johnstoni), described by Steindachner (1911) in the same work and apparently from the same
type locality as R. boulengeri. Consequently, the type locality may refer to Mount Ugoma (04.00000° S, 28.75000°
E, 2012 m elevation) in the southernmost part of the Itombwe Plateau, which is in the vicinity of Baraka (ca. 40 km)
and Kabambare in modern-day South Kivu and Maniema Provinces, eastern DRC.
Lectotype. Given the absence of a holotype, we designate the male specimen (NMW 16000-2) as the lectotype
for the species. This designation formally restricts the type locality to the Itombwe Plateau, DRC, as described
above. The specimens NMW 16000-1 and NMW 16000-3 thus become paralectotypes and no longer have any
name-bearing function.
FIGURE 7. Photographs of the lectotype and paralectotypes of Rhampholeon boulengeri. Adult male paralectotype (NMW
16000-1): (A)—lateral view; (B)—dorsal head view; (C)—lateral head view. Adult male lectotype (NMW 16000-2): (D)—
lateral view; (E)—dorsal head view; (F)—lateral head view. Adult female paralectotype (NMW 16000-3): (G)—lateral view;
(H)—dorsal head view; (I)—lateral head view. Photos by PW, Alice Schumacher, and Georg Gassner.

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464 · Zootaxa 5458 (4) © 2024 Magnolia Press
Referred specimens. UTEP 21715 (field no. EBG 1613) (Bichaka, 03.33872° S, 28.75595° E, 2100 m elevation),
UTEP 21716 (field no. EBG 1700) (Bichaka, 03.3374° S, 28.78804° E, 2311 m elevation), UTEP 21717–21718
(field nos. EBG 1701–1702) (Kisakala, 03.3373° S, 28.78791° E, 2264 m elevation), UTEP 22684–22686 (field nos.
DFH 4756–4758) (Mibengya, 03.46269° S, 28.58406° E, 2320 m elevation), UTEP 22687 (field no. DFH 4771)
(Elumbulumbu, 03.44536° S, 28.56246° E, 2470 m elevation), UTEP 22688 (field no. DFH 4779) (Elumbulumbu,
03.44501° S, 28.56195° E, 2464 m elevation), DRC, South Kivu Province, Itombwe Plateau (9 specimens).
Description of lectotype (NMW 16000-2). Adult male, SVL 51.3 mm and TL 16.4 mm. Body shape leaf-like.
Casque flattened, slightly elevated toward nape, with short head. Four elevated tubercles in center of casque. Neck
indistinct from head. Supra-orbital crests distinct with cluster of tubercles connected by ridge of tubercles between
crests. Rostral process 2.10 mm, composed of several elongated tubercles. Temporal crest discrete with five enlarged
tubercles extending posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised
tubercles, several raised higher above nares. Two raised tubercles below eye posteriorly and a third above rictus of
mouth. Body covered in nearly homogenous, flattened tubercles. Several larger conical tubercles present on dorsal
flanks around midbody. Largest body tubercle dorsal to forelimbs. Enlarged conical tubercles present on fore- and
hind limbs. Crenulated dorsal crest. Claws bicuspid.
Coloration of lectotype (in preservative). Photographs of the lectotype (NMW 16000-2) in preservative are
presented in Figure 7D–F. Head color is dark brown with lighter brown to pale orange coloration around mouth,
especially on lower jaw. Lighter brown hues begin behind the head and extend posteriorly and ventrally to just
beyond forelimbs, and dorsally along crest to about midbody. Forelimbs are darker brown with lighter color on
palms. Pale brown to white coloration is present on posterior third of body, extending from just beyond midbody to
cover hind limbs, rump, and tail. Larger tubercles on body and limbs dark brown to black, especially largest body
tubercle located dorsal to forelimbs.
TABLE 2. Meristic and mensural characters in type specimens of Rhampholeon boulengeri and summary of measurements
of examined specimens of topotypic and type material. Linear measurements (in mm) and scale counts are given. See
Materials and Methods for explanation of character abbreviations.
Types Summary morphometrics1
NMW 16000-1
Male
NMW 16000-2
Male
NMW 16000-3
Female
Males
n = 9
Females
n = 3
SVL 46.49 51.33 51.09 46.7 ± 4.9 (40.1–53.5) 45.8 ± 9.1 (35.3–51.1)
TL 12.95 16.43 12.67 15.6 ± 1.8 (12.9–17.6) 11.3 ± 2.1 (8.9–12.7)
TaL 59.44 67.67 63.76 62.3 ± 6.4 (53.4–71.1) 57.1 ± 11.1 (44.2–63.8)
HL 14.12 16.04 15.75 14.5 ± 0.9 (12.9–16.0) 14.2 ± 2.3 (11.6–15.8)
HW 7.98 8.57 8.89 8.2 ± 0.5 (7.3–8.9) 8.2 ± 1.2 (6.9–8.9)
HH 9.97 10.64 11.24 7.9 ± 1.4 (6.5–10.6) 8.1 ± 2.9 (5.7–11.2)
ML 7.85 9.66 9.32 9.3 ± 0.9 (7.9–10.5) 8.8 ± 1.3 (7.3–9.9)
CE 7.17 8.52 7.94 7.1 ± 0.6 (6.4–8.5) 6.9 ± 1.2 (5.6–7.9)
SL 4.77 5.05 6.29 5.0 ± 0.4 (4.3–5.5) 5.2 ± 1.1 (4.2–6.3)
ED 3.77 4.61 4.15 4.2 ± 0.3 (3.7–4.6) 4.0 ± 0.5 (3.5–4.5)
CC 6.87 7.21 7.34 6.6 ± 0.6 (5.8–7.5) 6.0 ± 1.2 (4.9–7.3)
IL 23.83 25.14 28.19 23.7 ± 2.8 (20.2–28.4) 23.6 ± 6.2 (16.5–28.2)
FLL 8.09 8.58 9.54 9.7 ± 0.9 (8.1–10.8) 8.7 ± 1.9 (6.6–9.9)
HLL 7.46 8.34 9.33 9.2 ± 0.9 (7.5–10.3) 8.2 ± 1.7 (6.2–9.3)
UL - - - 94 ± 6 (86–102)^ 101 ± 1.4 (100–102)^
LL - - - 91.7 ± 7.2 (80–102)^ 100 ± 0.0 (100–100)^
RP 1.58 2.1 1.68 1.5 ± 0.5 (0.9–2.2) 1.4 ± 0.4 (0.9–1.7)
TL/SVL 0.279 0.320 0.248 0.33 ± 0.03 (0.27–0.36) 0.24 ± 0.00 (0.24–0.25)
1Included in these data are measurements from the lectotype and paralectotypes. ^Sample sizes less than shown above.

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Variation. A summary of pairwise sequence divergence for three DNA markers (16S, ND2, and RAG-1) is
presented in Supplementary Material 1. A summary of morphometrics and measurements of the type specimens are
presented in Table 2 and comparative boxplots for the species in Figure 4. Photographs of the type specimens are
presented in Figure 7 and images displaying variation in life for the species are presented in Figure 8. Males and
females are similar in body size, but males have longer tails.
Distribution, natural history, and conservation. The species is distributed in montane forest in eastern DRC at
elevations between 2100 and 2470 m. One female specimen (UTEP 22684) collected on 9 June 2015 with an SVL
of 50.9 mm and TL of 12.3 mm was gravid. Most data on natural history previously assigned to the nominal taxon
(for a summary see Tilbury 2018) actually refers to other, cryptic species (see below).
Descriptions of five new species
Rhampholeon plumptrei sp. nov. Hughes, Behangana, Tilbury, Menegon, Kusamba, and Greenbaum
Plumptre’s pygmy chameleon
urn:lsid:zoobank.org:act:1A0EBA3F-A6DF-4A11-ABFB-8EF2880D47C8
Synonymy.
Rhampholeon boulengeri—de Witte 1965 (partim), Drewes & Vindum 1998, Nečas & Schmidt 2004 (partim), Matthee et al.
2004 (partim), Tolley et al. 2013 (partim), Stipala 2014, Glaw 2015 (partim), Tilbury & Tolley 2015 (partim), Spawls et al.
2018 (partim), Tilbury 2018 (partim)
Rhampholeon sp. 1—Hughes et al. 2018
Etymology. The specific epithet honors Andrew J. Plumptre for his efforts in promoting the conservation of Albertine
Rift biodiversity and whose leadership with the Wildlife Conservation Society has inspired scientists in Africa and
around the world.
Holotype. UTEP 22668 (field no. DFH 291), adult female, UGANDA, Western Region, Kigezi sub-region,
Kabale District, Bwindi Impenetrable National Park, Habinyanja Swamp, 00.98896° S, 29.63013° E, 1830 m
elevation, 14 June 2015, collected by D.F. Hughes and M. Behangana at night from forest vegetation about 1 m
above the ground (Fig. 9A).
Paratype (topotype). Same collection details as holotype, one adult female, UTEP 22669 (field no. DFH 292).
Paratypes. One sub-adult male, UTEP 22667 (field no. DFH 257), UGANDA, Western Region, Kigezi sub-
region, Kabale District, Bwindi Impenetrable National Park, Buhoma, 00.98918° S, 29.62944° E, 1846 m elevation,
14 June 2015, collected by E. Greenbaum and W. Rivera at night from forest vegetation. One small juvenile, UTEP
22666 (field no. DFH 252), UGANDA, Western Region, Kigezi sub-region, Kabale District, Bwindi Impenetrable
National Park, Buhoma, 00.99045° S, 29.61884° E, 1523 m elevation, 13 June 2015, collected by D.F. Hughes,
M. Behangana, E. Greenbaum, and W. Rivera at night from forest vegetation (Fig. 9B). Three juvenile males and
one adult female, UTEP 21685, 21386 (field nos. ELI 2764, 2816), UTEP 22757–22758 (field nos. ELI 2815, ELI
2817), UGANDA, Western Region, Kigezi sub-region, Kabale District, Bwindi Impenetrable National Park, Ihihizo
River, 00.99045° S, 29.61884° E, 1563 m elevation, 29 May 2014, collected by D.F. Hughes, K.A. Tolley, S. Davies,
A.A. Turner, and J. Kielgast at night from forest vegetation about 1–2 m above the ground (Figs. 9C, 9F).
Referred specimens. CAS 17682, 176864–176865, 176868, 176873, 176875–176876 (field nos. CAS-RCD
11445, 11474, 11476–11477; J. Vindum 0814, 0941, 0956; 01.00975° S, 29.62069° E, 1585 m elevation), 201681–
201682, 201687, 201689–201691 (field nos. J.V.V. 4146–4147, 4184, 4186–4187, 4334; 00.97833° S, 29.69500°
E, 1615 m elevation), UGANDA, Western Region, Bwindi Impenetrable National Park (13 specimens). PEM-R
16517 (field no. CT 355), UGANDA, Western Region, Kalinzu Central Forest Reserve (00.38583° S, 30.10638° E,
1468 m elevation) (1 specimen). UTEP 21686–21688, 22683 (field nos. CFS 1013w, 1025–1026w, 1050w), DRC,
North Kivu Province, Bunyantenge (00.26315° S, 28.94348° E, 1692 m elevation; Mount Vibende, 00.46247° S,
28.91721° E, 1892 m elevation; Kaunzo, 00.45250° S, 28.91518° E, 1675 m elevation; Kaluta River, 00.43317°
S, 28.90392° E, 1629 m elevation) (4 specimens). UTEP 21689–21690 (field nos. MUSE 10193, 10226), UTEP
22748–22752 (field nos. MUSE 10248–10252) (Mulwa; 02.37620° S, 28.32768° E, 1497 m elevation), UTEP
22753–22756 (field nos. MUSE 10259–10262) (Nyala; 02.34809° S, 28.28111° E, 1289 m elevation), MTSN

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6898–6899 (Madiriri; 02.30926° S, 28.64935° E, 2036 m elevation), DRC, South Kivu Province, Kahuzi-Biega
National Park (13 specimens).
Diagnosis. Rhampholeon plumptrei sp. nov. is in the subgenus Rhinodigitum because of its distinctly bicuspid
claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily distinguishing
it from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae Broadley,
1971, R. marshalli Boulenger, 1906, R. spectrum (Buchholz, 1874), R. spinosus (Matschie, 1892) R. temporalis
(Matschie, 1892), and R. viridis Mariaux & Tilbury, 2006. Rhampholeon plumptrei sp. nov. can be distinguished
from all other Rhampholeon species by the following combination of traits: (1) lack of prominent mite pockets in
the inguinal region distinguishes it from R. beraduccii Mariaux & Tilbury, 2006, R. platyceps Günther, 1893, R.
chapmanorum Tilbury, 1992, R. maspictus Branch, Bayliss & Tolley, 2014, R. tilburyi Branch, Bayliss & Tolley,
2014, R. bruessoworum Branch, Bayliss & Tolley, 2014, and R. nebulauctor Branch, Bayliss & Tolley, 2014; (2)
presence of prominent mite pockets in the axillary region distinguishes it from R. nchisiensis (Loveridge, 1953)
and R. acuminatus Mariaux & Tilbury, 2006; (3) distinct supra-orbital and canthal crests distinguishes it from
R. hattinghi Tilbury & Tolley, 2015; (4) geographic restriction in the Albertine Rift, central Uganda, and eastern
Kenya distinguishes it from R. uluguruensis Tilbury & Emmrich, 1996, R. moyeri Menegon, Salvidio & Tilbury,
2002, R. colemani Menegon et al., 2022, R. sabini Menegon et al., 2022, R. rubeho Menegon et al., 2022, R. nicolai
Menegon et al., 2022, R. waynelotteri Menegon et al., 2022, and R. princeeai Menegon et al., 2022; (5) slightly
shorter tails and shorter cranial crest gaps in males distinguishes it from R. boulengeri; (6) slightly longer inter-limb
lengths in males, larger eye diameters and longer inter-limb length in females, and longer snout lengths in both sexes
distinguishes it from R. monteslunae sp. nov.; (7) slightly shorter head, snout, and mouth lengths in females, and
longer inter-limb length in females distinguishes it from R. nalubaale sp. nov.; (8) genetic divergence and non-
overlapping elevational range at sites of co-occurrence distinguishes it from R. bombayi sp. nov.; and (9) genetic
divergence and allopatric distribution distinguishes it from R. msitugrabensis sp. nov.
Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers
(16S, ND2, and RAG-1) among individuals of R. plumptrei sp. nov. and other Rhampholeon species is presented in
Supplementary Material 1.
Description of holotype (UTEP 22668). Adult female, SVL 49.2 mm and TL 12.5 mm. Body shape leaf-like.
Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests distinct with cluster of tubercles
connected by a ridge with 16 tubercles across casque and 22 tubercles from peak-to-peak of crests. Rostral process
1.63 mm, composed of elongated tubercles. Temporal crest discrete with several enlarged tubercles extending
posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised tubercles, one
raised higher than others near snout. Ninety-eight upper and 90 lower labial tubercles are present along tip of snout
to rictus of mouth. Body covered in nearly homogenous, flattened tubercles. Several larger conical tubercles present
on dorsal flanks around midbody. Crenulated dorsal crest, more prominent from mid-body to nape. Many enlarged
conical tubercles present on limbs. Claws markedly bicuspid.
Coloration of holotype (in life). A photograph of the holotype (in life) is presented in Figure 9A. Background
color is orangish brown with more white coloration towards ventral area. Head is similar in color to body. Gular
region is white, and this color extends onto ventral region. Two diagonal dark brown lines extend from near dorsal
crest posteriorly to lateral flanks, resembling veins of a leaf. Tail is similar color as body with obvious white
coloration underneath. Several small greenish to yellow patches present on upper limbs and near nape. Larger body
tubercles are darker brown.
Variation. A summary of descriptive morphometrics for R. plumptrei sp. nov. is presented in Table 3, comparative
boxplots in Figure 4, and measurements of the type specimens in Table 5. Photographs displaying color variation in
life are presented in Figure 9. Morphological proportions are generally consistent with those of the holotype. Males
have smaller body sizes (males [M]: mean 42.5 mm, range 27.7–53.3 mm, n = 18; females [F]: mean 50.8 mm,
range 30.3–60.2 mm, n = 18) and longer tails than females (M: mean 12.3 mm, range 9.0–15.7 mm, n = 18; F: mean
11.9 mm, range 8.3–13.9 mm, n = 18). Body coloration is consistently brown, gray, or tan, occasionally with reddish
or white hues. Often exhibits one, sometimes two, dark brown lines on the lateral flanks extending diagonally from
the dorsal crest toward the hind limbs, resembling veins of a leaf. In juveniles, the top of the head and legs can be
black. Usually, the largest conical tubercle near the nape is much darker than the body. Legs and tail can be a darker
brown than the body.
Reproduction. Five females were gravid (UTEP 21689, UTEP 22748, UTEP 22750, UTEP 22755, and UTEP
21686). Gravid females had a mean SVL of 55.4 mm (49.2–60.2 mm) and a mean TL of 12.4 mm (10.9–13.9 mm).

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These females were collected in April (n = 1), July (n = 1), and November (n = 3). The smallest specimen examined
(UTEP 22666) was collected on 13 June 2015 with SVL 22.2 and TL 5.8 from Bwindi Impenetrable National Park,
Uganda.
Distribution, natural history, and conservation. Rhampholeon plumptrei sp. nov. is found in montane and
sub-montane forests at an elevation range from 1203–2269 m. This species has a very large geographic distribution
from eastern DRC (Kahuzi-Biega National Park) to western Kenya (Kakamega Forest National Reserve), including
its occurrence in several protected areas (e.g., Bwindi Impenetrable National Park, Uganda) (Hughes et al. 2018).
We note there is also a documented population in Mabira Central Forest Reserve, Uganda, found at the Rainforest
Lodge (00.38202° N, 33.01702° E, 1325 m), which represents an intermediate site between the Albertine Rift and
FIGURE 8. Photographs of various individuals of Rhampholeon boulengeri in life. (A)—Adult male (UTEP 21715) from the
vicinity of Bichaka, DRC; (B)—Uncollected adult male from the Itombwe Plateau, DRC; (C, D)—Adult male (UTEP 21716)
from the vicinity of Bichaka, DRC; (E)—Habitat view of the forest at Bichaka, DRC; (F)—Another habitat view of the forest
at Bichaka, DRC.

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FIGURE 9. Photographs of various individuals of Rhampholeon plumptrei sp. nov. in life. (A)—Adult female holotype (UTEP
22668) from Bwindi Impenetrable National Park, Uganda; (B)—Juvenile (UTEP 22666) from Bwindi Impenetrable National
Park, Uganda; (C)—Male paratype (UTEP 21685) from Bwindi Impenetrable National Park, Uganda; (D)—Female (UTEP
22749) from Kahuzi-Biega National Park, DRC; (E)—Female (UTEP 22750) from Kahuzi-Biega National Park, DRC; (F)—
Female (UTEP 21386) from Bwindi Impenetrable National Park, Uganda; (G)—Habitat view of the type locality showing the
distinction between the protected forest and agriculture; (H)—Habitat view of a swamp in the type locality showing an area
known as “the neck.”

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the easternmost populations in Kenya (MB, pers. obs). At the type locality, Drewes & Vindum (1998) indicated that
this species was found at elevations up to 2100 m where it was seen perched from 100 to 500 mm above the ground
on ferns along trails and near streams. Furthermore, Drewes & Vindum (1998) stated that this species was the most
common lizard that they encountered during their night surveys and that females had a maximum clutch size of four
eggs. In contrast, extensive mark-recapture chameleon surveys across five nights in November 2018 by one of us
(DFH) in Bwindi Impenetrable National Park along the Hamufa Trail (near Ruhija) detected only a single female
R. plumptrei sp. nov. at an elevation of 2269 m, which was sleeping on a shrub at a perch height of 305 mm and a
perch diameter of 1.14 mm. During five nights with three surveyors, this chameleon species was actually the least
frequently encountered compared with Trioceros johnstoni (> 70 captures) and Kinyongia tolleyae (> 20 captures).
The surveys of Drewes & Vindum (1998) found only a single specimen at 2100 m elevation (Ntengere River) with
nearly all of their collections coming from mid-elevation sites around Buhoma, which occurs at about 1500 m
elevation. In this study, only two specimens that we examined were found above 2000 m elevation, and just four
specimens were from 1700 to 2000 m elevation. Taken together, it seems that R. plumptrei sp. nov. is most common
at mid-elevations (1200–1700 m), uncommon at higher elevations (1700–2000 m), and rare above 2000 m. For a
detailed list of lizard species present at the type locality see Drewes & Vindum (1998), and for Kakamega Forest,
Kenya, see Wagner & Böhme (2007) and Lötters et al. (2007).
TABLE 3. Summary of meristic and mensural characters in specimens of Rhampholeon plumptrei sp. nov., R. nalubaale
sp. nov., and R. bombayi sp. nov. Linear measurements (in mm) and scale counts are given as mean ± standard deviation,
followed by range in parentheses. See Materials and Methods for explanation of character abbreviations.
Rhampholeon plumptrei sp. nov. Rhampholeon nalubaale sp. nov. Rhampholeon bombayi sp. nov.
Males
n = 18
Females
n = 18
Females
n = 19
Males
n = 14
Females
n = 22
SVL 42.5 ± 7.3 (27.7–53.3) 50.8 ± 6.3 (30.3–60.2) 49.4 ± 5.2 (39.3–56.2) 41.5 ± 5.3 (31.0–47.6) 47.9 ± 5.2 (40.8–59.1)
TL 12.3 ± 2.2 (9–15.7) 11.9 ± 1.4 (8.3–13.9) 11.9 ± 1.3 (8.8–13.9) 12.9 ± 2.8 (9.3–18.9) 11.4 ± 0.9 (10.0–13.4)
ToL 54.8 ± 8.9 (37.5–68.4) 62.7 ± 7.2 (38.6–72.6) 61.4 ± 6.3 (48.1–69.5) 54.4 ± 7.4 (40.3–63.9) 59.3 ± 5.2 (51.7–70.1)
HL 13.4 ± 1.7 (10.6–16.3) 15.1 ± 1.3 (11.5–16.6) 15.2 ± 1.3 (12.3–16.9) 13.5 ± 1.5 (10.6–15.1) 14.6 ± 0.9 (13.0–16.9)
HW 7.7 ± 1.1 (5.6–9.6) 8.6 ± 0.9 (5.6–9.6) 8.2 ± 0.7 (6.6–9.4) 7.7 ± 0.7 (6.3–8.7) 8.2 ± 0.5 (7.6–9.5)
HH 7.1 ± 0.7 (5.8–8.3) 7.8 ± 0.7 (6.5–10.3) 7.7 ± 0.8 (5.6–8.9) 7.0 ± 0.6 (5.9–7.9) 7.5 ± 0.5 (6.2–8.5)
ML 8.5 ± 1.2 (6.3–10.9) 9.8 ± 0.9 (8.3–11.8) 9.9 ± 1.1 (7.8–11.6) 8.6 ± 1.0 (6.2–10.0) 9.5 ± 0.8 (8.2–11.6)
CE 6.5 ± 0.9 (5.1–7.9) 7.1 ± 0.5 (5.6–8.0) 7.2 ± 0.6 (5.7–8.1) 6.6 ± 0.7 (5.3–7.5) 6.9 ± 0.4 (6.4–7.9)
SL 4.7 ± 0.7 (3.4–5.9) 5.3 ± 0.5 (3.9–6.2) 5.4 ± 0.5 (4.2–6.3) 4.5 ± 0.5 (3.7–5.1) 5.2 ± 0.6 (4.2–7.2)
ED 4.1 ± 0.7 (2.7–5.6) 4.4 ± 0.5 (3.4–5.3) 4.5 ± 0.4 (3.4–5.2) 4.1 ± 0.6 (3.1–4.9) 4.2 ± 0.3 (3.8–5.0)
CC 5.8 ± 0.8 (3.9–6.9) 6.0 ± 0.5 (4.9–6.9) 5.7 ± 0.6 (4.5–6.6) 5.6 ± 0.8 (3.9–6.9) 5.7 ± 0.5 (4.8–6.7)
IL 22.1 ± 4.5 (11.6–28.0) 28.6 ± 3.7 (17.1–33.8) 27.3 ± 3.4 (20.0–31.3) 21.3 ± 2.7 (15.8–24.7) 25.8 ± 3.2 (21.1–32.4)
FLL 8.9 ± 1.4 (6.3–10.9) 9.8 ± 0.8 (7.8–11.0) 9.9 ± 1.4 (6.9–12.7) 8.8 ± 1.2 (6.4–10.7) 9.6 ± 0.7 (8.8–11.3)
HLL 8.4 ± 1.5 (5.7–10.6) 9.3 ± 0.9 (6.5–10.8) 9.3 ± 1.1 (6.6–11.0) 8.4 ± 1.3 (5.9–10.3) 9.0 ± 0.7 (8.1–10.7)
UL 93.1 ± 5.9 (82–101) 96.6 ± 8.4 (80–110) 92.5 ± 5.9 (76–102) 94.3 ± 6.2 (88–106) 94.8 ± 8.3 (78–108)
LL 92.8 ± 5.8 (84–104) 93.8 ± 5.7 (84–104) 92 ± 4.1 (86–98) 89.3 ± 7.8 (72–100) 91.6 ± 6.3 (80–106)
RP 1.4 ± 0.4 (0.9–2.1) 1.4 ± 0.2 (0.9–1.8) 1.5 ± 0.2 (1.0–1.9) 1.4 ± 0.4 (0.9–2.0) 1.4 ± 0.3 (0.9–1.9)
TL/SVL 0.29 ± 0.04 (0.20–0.35) 0.23 ± 0.03 (0.20–0.28) 0.24 ± 0.02 (0.21–0.27) 0.31 ± 0.05 (0.24–0.44) 0.24 ± 0.03 (0.18–0.30)
Rhampholeon nalubaale sp. nov. Hughes, Behangana, Lukwago, Kusamba, and Greenbaum
Goddess pygmy chameleon
urn:lsid:zoobank.org:act:F997E4A5-298C-4137-9528-BF9A4DD04E3A
Synonymy.
Rhampholeon boulengeri—Schmidt 1919, de Witte 1965 (partim), Vonesh 2001, Kvift 2011, Tolley et al. 2013 (partim), Tilbury
& Tolley 2015 (partim), Spawls et al. 2018 (partim), Tilbury 2018 (partim)
Rhampholeon sp. 2—Hughes et al. 2018

HUGHES ET AL.
470 · Zootaxa 5458 (4) © 2024 Magnolia Press
Etymology. The specific epithet is derived from the Luganda word, nalubaale, which means a spirit of feminine
qualities (i.e., goddess), and is also the name for Lake Victoria (Kenny 1977).
Holotype. UTEP 22673 (field no. DFH 585), adult female, UGANDA, Western Region, Bunyoro sub-region,
Masindi District, Budongo Central Forest Reserve, 01.72992° N, 31.55535° E, 1124 m elevation, 2 July 2015,
collected by D.F. Hughes, W. Lukwago, and M. Behangana at night from forest vegetation about 1 m above the
ground (Fig. 10A).
Paratype (topotype). Same collection details as holotype, two adult females, UTEP 22671–22672 (field nos.
DFH 583–584) (Fig. 10B–C).
Paratypes. Three adult females, UTEP 21693–21694, 22670 (field nos. DFH 558–560), UGANDA, Western
Region, Bunyoro sub-region, Masindi District, Budongo Central Forest Reserve, Sonso River, 01.71885° N,
31.54095° E, 1073 m elevation, 1 July 2015, collected by D.F. Hughes and W. Lukwago at night from forest
vegetation about 1 m above the ground (Fig. 10D).
Referred specimens. UTEP 21695–21697 (field nos. ELI 464–465, EBG 2737), UTEP 22744–22747 (field
nos. ELI 466, 479–481), DRC, South Kivu Province, Bizombo (03.02880° S, 28.28243° E, 1132 m elevation) (7
specimens) (Fig. 10F). UTEP 21700 (field no. DFH 1101) and CAS 204371 (field no. JRV 450), UGANDA, Western
Region, Kibale National Park, Ngogo, 00.49795° N, 30.42262° E, 1363 m elevation (UTEP 21700 [Fig. 10E]), and
Kanyawara, 00.56666° N, 30.35000° E, 1506 m elevation (CAS 204371) (2 specimens). UTEP 21691–21692 (field
nos. CRSN HERP 2833, 2837), DRC, Ituri Province, Bongobongo, 02.19766° N, 30.11468° E, 1258 m elevation
(2 specimens). UTEP 21698–21699 (field nos. MUSE 10178, 10589), DRC, South Kivu Province, Kahuzi-Biega
National Park, Ikundwe, 02.57567° S, 28.16142° E, 1137 m elevation (UTEP 21698), and Kyasa, 01.75368° S,
27.61806° E, 975 m elevation (UTEP 21699) (2 specimens).
Diagnosis. Rhampholeon nalubaale sp. nov. is in the subgenus Rhinodigitum because of its distinctly bicuspid
claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily distinguishing it
from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae, R. marshalli, R.
spectrum, R. spinosus, R. temporalis, and R. viridis. Rhampholeon nalubaale sp. nov. can be distinguished from
all other Rhampholeon species by the following combination of traits: (1) lack of prominent mite pockets in the
inguinal region distinguishes it from R. beraduccii, R. platyceps, R. chapmanorum, R. maspictus, R. tilburyi, R.
bruessoworum, and R. nebulauctor; (2) presence of prominent mite pockets in the axillary region distinguishes it
from R. nchisiensis and R. acuminatus; (3) distinct supra-orbital and canthal crests distinguishes it from R. hattinghi;
(4) geographic restriction in the Albertine Rift distinguishes it from R. uluguruensis, R. moyeri, R. colemani, R.
sabini, R. rubeho, R. nicolai, R. waynelotteri, and R. princeeai; (5) larger inter-limb length in females, genetic
divergence, and non-overlapping elevational range at parapatric sites distinguishes it from R. boulengeri; (6) larger
mean body size, larger eye diameters, longer mouth, snout, inter-limb, and hind limb lengths in females distinguishes
it from R. monteslunae sp. nov.; (7) slightly longer head, snout, and mouth lengths, and shorter inter-limb length in
females distinguishes it from R. plumptrei sp. nov.; (8) slightly longer head, snout, and mouth lengths, larger eye
diameters, and longer inter-limb and hind limb lengths in females distinguishes it from R. bombayi sp. nov.; (9)
longer inter-limb, hind limb, and head lengths in females distinguishes it from R. msitugrabensis sp. nov.
Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers
(16S, ND2, and RAG-1) among individuals of R. nalubaale sp. nov. and other Rhampholeon species is presented in
Supplementary Material 1.
Description of holotype (UTEP 22673). Adult female, SVL 48.9 mm and TL 10.8 mm. Gravid (see Reproduction
below). Body shape leaf-like. Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests
distinct with cluster of tubercles connected by a ridge with 14 tubercles across casque and 19 tubercles from peak-
to-peak of crests. Rostral process 1.78 mm, composed of elongated tubercles. Temporal crest discrete with several
enlarged tubercles extending posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists
of raised tubercles, one raised higher than others near snout. Ninety-eight upper and 94 lower labial tubercles
present along tip of snout to rictus of mouth. Body covered in nearly homogenous, flattened tubercles. Crenulated
dorsal crest, more prominent from mid-body to nape. Many enlarged conical tubercles present on limbs, tail, and
lower dorsal flanks. Claws markedly bicuspid.
Coloration of holotype (in life). A photograph of the holotype is presented in Figure 10A. Background color tan
with a yellow-orange hue. Top of head a whitish coloration. Head generally light tan and indistinct from body. Gular
region white, which extends onto ventral area. One diagonal dark red line extends from near dorsal crest posteriorly

FIVE NEW RHAMPHOLEON FROM CENTRAL AFRICA Zootaxa 5458 (4) © 2024 Magnolia Press · 471
FIGURE 10. Photographs of various individuals of Rhampholeon nalubaale sp. nov. in life. (A)—Adult female holotype
(UTEP 22673) from Budongo Central Forest Reserve, Uganda; (B)—Adult female paratopotype (UTEP 22671) with recently
laid eggs from Budongo Central Forest Reserve, Uganda; (C)—Adult female paratopotype (UTEP 22672) from Budongo
Central Forest Reserve, Uganda; (D)—Adult female paratype (UTEP 21694) from Budongo Central Forest Reserve, Uganda;
(E)—Adult female (UTEP 21700) from Kibale Forest National Park, Uganda; (F)—Adult female (UTEP 21695) from Bizombo,
DRC; (G)—Habitat view of the forest at Bizombo, DRC; (H)—Another habitat view of the forest at Bizombo, DRC.

HUGHES ET AL.
472 · Zootaxa 5458 (4) © 2024 Magnolia Press
TABLE 4. Summary of meristic and mensural characters in specimens of Rhampholeon msitugrabensis sp. nov. and R.
monteslunae sp. nov. Linear measurements (in mm) and scale counts are given as mean ± standard deviation, followed
by range in parentheses; see Materials and Methods for explanation of character abbreviations.
Rhampholeon msitugrabensis sp. nov. Rhampholeon monteslunae sp. nov.
Males
n = 3
Females
n = 4
Males
n = 8
Females
n = 10
SVL 41.7 ± 7.1 (33.6–46.8) 47.8 ± 1.8 (45.7–49.4) 41.1 ± 5.5 (33.4–50.9) 40.4 ± 9.2 (30.4–54.4)
TL 12.4 ± 1.1 (11.5–13.7) 10.9 ± 0.4 (10.5–11.3) 13.4 ± 1.5 (11.2–15.6) 10.9 ± 1.6 (8.2–13.7)
TaL 54.0 ± 7.9 (45.1–60.5) 58.8 ± 2.2 (56.1–60.6) 54.4 ± 6.6 (44.7–66.5) 51.4 ± 10.3 (38.7–68.1)
HL 13.8 ± 1.5 (12.1–14.8) 14.3 ± 0.3 (14.0–14.5) 13.8 ± 1.3 (11.9–15.7) 13.6 ± 1.9 (11.1–16.5)
HW 8.1 ± 0.8 (7.1–8.6) 8.3 ± 0.3 (7.9–8.7) 7.9 ± 0.7 (7.2–9.1) 7.7 ± 1.4 (5.6–9.5)
HH 7.4 ± 0.8 (6.8–8.3) 7.4 ± 0.6 (6.8–7.9) 7.5 ± 0.6 (6.8–8.3) 7.2 ± 0.9 (5.8–9.2)
ML 8.9 ± 0.6 (8.2–9.4) 9.6 ± 0.2 (9.3–9.7) 8.7 ± 0.7 (7.8–9.7) 8.6 ± 0.9 (6.9–10.3)
CE 7.1 ± 0.9 (6.1–7.8) 6.9 ± 0.2 (6.7–7.2) 6.9 ± 0.9 (5.9–8.4) 7.1 ± 1.3 (5.8–9.9)
SL 4.7 ± 0.7 (4.1–5.4) 4.8 ± 0.3 (4.5–5.1) 4.4 ± 0.5 (3.7–4.9) 4.6 ± 0.6 (3.9–5.4)
ED 3.9 ± 0.3 (3.8–4.3) 3.9 ± 0.3 (3.5–4.2) 3.9 ± 0.4 (3.5–4.8) 3.8 ± 0.6 (3.1–4.8)
CC 6.1 ± 0.7 (5.4–6.8) 5.5 ± 0.3 (5.2–5.8) 5.7 ± 0.8 (4.6–7.1) 5.8 ± 1.1 (4.1–7.5)
IL 21.5 ± 4.2 (16.8–24.8) 25.8 ± 0.3 (25.5–26.2) 20.6 ± 3.1 (16.5–25.1) 21.3 ± 7.5 (13.2–33.4)
FLL 8.8 ± 1.0 (8.0–9.9) 9.9 ± 0.2 (9.8–10.1) 9.6 ± 1.0 (8.3–11.3) 9.1 ± 1.7 (6.9–11.7)
HLL 8.4 ± 1.2 (7.0–9.4) 9.2 ± 0.5 (8.8–9.8) 8.9 ± 0.9 (7.6–10.3) 8.2 ± 1.9 (5.8–10.9)
UL 88.7 ± 6.4 (84–96) 93 ± 3.5 (88–96) 88.5 ± 7.1 (76–100) 93.8 ± 10.2 (80–108)
LL 86 ± 3.5 (82–88) 89.5 ± 4.4 (86–96) 84 ± 5.3 (74–90) 89.2 ± 7.7 (76–100)
RP 1.3 ± 0.3 (1.0–1.6) 1.0 ± 0.2 (0.8–1.3) 1.3 ± 0.4 (0.8–1.8) 1.3 ± 0.5 (0.7–2.4)
TL/SVL 0.30 ± 0.04 (0.26–0.34) 0.22 ± 0.00 (0.22–0.23) 0.32 ± 0.03 (0.29–0.38) 0.27 ± 0.04 (0.19–0.33)
TABLE 5. Meristic and mensural characters in type specimens of Rhampholeon plumptrei sp. nov. Linear measurements
(in mm) and scale counts are given. See Materials and Methods for explanation of character abbreviations.
UTEP
22668
Female
Holotype
UTEP
22669
Female
Paratopotype
UTEP
22667
Male
Paratype
UTEP
21685
Male
Paratype
UTEP
22757
Male
Paratype
UTEP
22758
Male
Paratype
UTEP
22666*
Juvenile
Paratype
UTEP
21386**
Female
Paratype
SVL 49.22 46.23 31.53 33.72 27.71 29.75 22.19 48
TL 12.46 10.95 10.31 10.51 9.83 9.41 5.82 13
TaL 61.68 57.18 41.84 44.23 37.54 39.16 28.01 61
HL 14.69 13.68 10.67 11.94 10.56 10.61 - -
HW 8.32 7.66 5.95 6.73 5.77 5.57 - -
HH 7.94 7.49 5.77 6.96 5.96 6.32 - -
ML 9.97 8.25 6.25 7.9 6.94 6.39 - -
CE 7.29 6.67 5.24 5.68 5.36 5.13 - -
SL 4.92 5.08 3.85 4.01 3.64 3.37 - -
ED 4.44 3.7 3.3 3.53 3.08 2.7 - -
CC 5.87 5.99 3.92 4.97 4.92 4.83 - -
IL 28.81 26.59 16.28 17.39 11.63 14.42 - -
FLL 9.66 9.03 6.28 7.59 6.81 6.66 - -
HLL 8.79 8.45 5.81 6.79 6.06 5.65 - -
UL 98 96 90 92 88 84 - -
LL 90 88 92 94 88 92 - -
RP 1.63 1.77 1.15 1.14 1.14 1.24 - -
TL/SVL 0.253 0.237 0.327 0.312 0.355 0.316 0.262 0.271
*Specimen was a recently hatched juvenile. **Specimen measured using a standard ruler because it was used in the
neuroanatomical study of Hughes et al. (2016).

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to lateral flank with second line posterior, but faint. Tail a similar tan color as body. Several large tubercles on body
and limbs white, especially towards ventral area. Largest body tubercle near neck dark brown. Limbs slightly darker
brown color than body. Several light orange and yellow patches on body, especially along dorsal crest and near
forelimb-nape region.
Variation. A summary of descriptive morphometrics for R. nalubaale sp. nov. is presented in Table 3, comparative
boxplots in Figure 4, and measurements of the type specimens in Table 6. Photographs displaying color variation
in life are presented in Figure 10. Morphological proportions are generally consistent with those of the holotype.
Only females have been found. Relatively large species (mean SVL 49.4 mm, range 39.3–56.2 mm, n = 19) with a
prominent rostral process (mean 1.5 mm, range 1.0–1.9 mm, n = 19). Body coloration is consistently brown to tan,
usually with red-orange hues. Most exhibited one, sometimes two, dark brown to red lines on the lateral flanks that
extended diagonally from the dorsal crest toward the hind limbs, resembling veins of a leaf.
Reproduction. Five females were gravid (UTEP 21691, UTEP 21692, UTEP 21698, UTEP 22672, and UTEP
22673 [holotype]). Gravid females had a mean SVL of 52.9 mm (48.9–56.2 mm) and a mean TL of 12.2 mm (10.8–
13.8 mm). These females were collected in January (n = 2) and July (n = 3). One female had a clutch size of 3 eggs
(Fig. 10B). The smallest specimen examined (UTEP 21694) was collected on 1 July 2015 with SVL 39.3 mm and
TL 10.8 mm from Budongo Central Forest Reserve, Uganda.
Distribution, natural history, and conservation. Rhampholeon nalubaale sp. nov. is found in sub-montane
forests at an elevation range from 513–1506 m. Specimens have been collected from Kibale National Park and
Budongo Central Forest Reserve in Uganda, and Kahuzi-Biega National Park and Itombwe Natural Reserve in
DRC. Vonesh (2001) recorded this species from Kibale National Park, where he found it to be the most commonly
observed chameleon species relative to T. ituriensis and K. tolleyae, and was most often found on low shrubs.
Extensive mark-recapture chameleon surveys in October 2018 by one of us (DFH) in Kibale National Park near
Kanyawara Research Station across seven nights found just four females sleeping at a mean perch height of 375
mm (range 254–508 mm) and a mean perch diameter of 2.94 mm (range 1.79–4.67 mm). One female was gravid,
and no individuals were recaptured across the seven consecutive survey nights. In fact, T. ituriensis was found to
be much more common than R. nalubaale sp. nov. during those surveys with 33 individuals captured 42 times
compared to the four individuals captured each once for the pygmy chameleon. We examined a specimen (UTEP
22673) under ultraviolet light and found that its facial tubercles fluoresced blue (Fig. 11), indicating that they
are of bony origin. Prötzel et al. (2018) found that this phenomenon of bone fluorescence is actually widespread
in Chamaeleonidae, especially in species that live in forested habitats with relatively higher levels of ambient
ultraviolet light. It is currently thought that chameleons that possess this trait have co-opted bone fluorescence for
signaling between individuals (Prötzel et al. 2018). We note that this is the first published account of fluorescence in
the genus Rhampholeon, and while we did not examine other species under ultraviolet light, this has been observed
in several other species of Rhampholeon (K.A. Tolley, pers. comm.); thus, we anticipate that most species in the
genus will also exhibit this trait. The current range of R. nalubaale sp. nov. falls within the boundaries of at least
two national parks and we suspect it may be present in other protected areas with suitable lowland forest habitat
across the region. Budongo Central Forest Reserve and Itombwe Natural Reserve are also protected, but subjected
to greater anthropogenic pressure relative to the national parks because they both allow some level of human use
of the forests (Kujirakwinja et al. 2018; Lukwago et al. 2020) and law enforcement in the latter reserve is weak
(Greenbaum & Kusamba 2012; EG, pers. obs.).
Schmidt (1919) inspected a large collection of R. boulengeri from the Ituri Forest, DRC, housed at the American
Museum of Natural History (AMNH), New York, which he noted were “all, curiously, females.” The collection
included 61 specimens from Medje (Haut-Uele Province) and two specimens from Poko (Bas-Uele Province), both
low-elevation villages in DRC. Based on Schmidt’s (1919) collection, Hall (1970) speculated that R. boulengeri
(sensu lato) was a possible case of parthenogenesis in Chamaeleonidae, and to corroborate, he examined specimens
at the Museum of Comparative Zoology (MCZ), Harvard. Hall (1970) examined at least four specimens (MCZ
Herp R-53983–53986) from a different site in the Ituri Forest (Mayala [513 m elevation]) collected by C.J.P. Ionides
and, by exposing their gonads, he found that these specimens were all females. We also concluded that these four
specimens were females based recent photographs taken by the current collections manager at the MCZ, who
likewise indicated that they are all females based on visual inspections of the specimens (S. Kennedy-Gold, pers.
comm.). Hall (1970) also discussed other specimens in the MCZ collected from Central Africa that included males,
such as those from Ja River and Kribi, Cameroon, and also from Idjwi Island, DRC. The Cameroonian specimens

HUGHES ET AL.
474 · Zootaxa 5458 (4) © 2024 Magnolia Press
cannot be R. boulengeri because it is now restricted to the Albertine Rift, and thus they would be R. spectrum.
The three Idjwi Island specimens (MCZ Herp R-47297–47299) were collected by Arthur Loveridge in 1939 and
included at least one male (48 mm SVL and 16 mm TL) and one gravid female (47 mm SVL and 14 mm TL with 3
eggs) (Loveridge 1942). A photograph of one of these specimens is provided by Loveridge (1942: Plate 5, Figure 1),
which is clearly a male based on its long tail with an obvious hemipenal bulge. According to Loveridge (1944), the
specimens collected on Idjwi Island were taken from near the Upper Mulinga River around 1981 m elevation. Based
on the elevation at the collecting site and body sizes presented in Loveridge (1942), these Idjwi Island specimens
would likely represent a different member of the R. boulengeri complex, probably R. bombayi sp. nov., as this
species occurs in the region at other high-elevation sites where numerous males have been collected. Ultimately,
Hall (1970) did not rule out parthenogenesis in R. boulengeri (sensu lato) because males possessed “greatly enlarged
hemipenal swellings and much longer tails,” and thus he deemed sex-determination errors by Schmidt (1919) highly
unlikely. Later, Kearney et al. (2009) dismissed the possibility of parthenogenesis in R. boulengeri (sensu lato)
because the evidence of a sex-ratio bias was considered too weak, and no further evidence had come to light since
Hall (1970).
Our findings for R. nalubaale sp. nov., where only females have been in collections from numerous lowland
sites across the region (see Fig. 2; Table 3), nonetheless, extends Schmidt’s (1919) observation from the villages
of Medje (715 m elevation) and Poko (634 m elevation). In contrast, at least two males have been found for all
other species of the complex (see Tables 1–2), and for some species, just as many males have been collected as
females, such as R. plumptrei sp. nov. Moreover, the level of intraspecific genetic divergence among samples of R.
nalubaale sp. nov. was negligible to non-existent, even across vast geographic scales (some samples are separated
by > 600 km straight-line distance), and was much lower than the ranges among samples of any other member of
the R. boulengeri complex (excluding R. boulengeri [sensu stricto] with only four samples that have been sequenced
from the same time and place). For example, using the most variable locus of Hughes et al. (2018), ND2, the range
of intraspecific variation for R. nalubaale sp. nov. was 0.1–0.3% (n = 10), whereas the ranges for the other new
species were much larger, especially for those species with comparably large geographic ranges: R. plumptrei sp.
nov. (0.1–3.4%, n = 14), R. bombayi sp. nov. (0.1–4.4%, n = 12), R. msitugrabensis sp. nov. (2%, n = 2), and R.
monteslunae sp. nov. (0.1–1.1%, n = 7).
TABLE 6. Meristic and mensural characters in type specimens of Rhampholeon nalubaale sp. nov. Linear measurements
(in mm) and scale counts are given. See Materials and Methods for explanation of character abbreviations.
UTEP 22673
Female
Holotype
UTEP 22671
Female
Paratopotype
UTEP 22672
Female
Paratopotype
UTEP 21693
Female
Paratype
UTEP 21694
Female
Paratype
UTEP 22670
Female
Paratype
SVL 48.94 51.48 52.56 52.98 39.25 39.54
TL 10.78 12.57 11.61 13.87 10.84 10.91
TaL 59.72 64.05 64.17 66.85 50.09 50.45
HL 14.91 16.05 15.16 16.82 13.03 12.99
HW 8.1 8.56 8.23 8.43 7.34 7.11
HH 7.68 8.49 8.08 8.1 7.59 6.3
ML 9.1 10 9.88 10.81 8.13 8.12
CE 7.16 7.26 6.95 7.65 6.28 6.17
SL 5 5.55 5.26 5.61 4.76 4.53
ED 4.39 4.45 4.51 4.68 4.01 3.89
CC 6.17 6.56 4.99 6.28 5.44 5.15
IL 28.6 27.84 29.98 29.83 20.85 20.95
FLL 9.78 10.3 9.59 10.36 7.53 7.47
HLL 9.35 9.72 9.49 9.49 7.46 7.24
UL 98 96 96 94 98 102
LL 94 98 96 88 90 94
RP 1.78 1.53 1.92 1.65 1.78 1.7
TL/SVL 0.220 0.244 0.221 0.262 0.276 0.276

FIVE NEW RHAMPHOLEON FROM CENTRAL AFRICA Zootaxa 5458 (4) © 2024 Magnolia Press · 475
FIGURE 11. Photographs illustrating bone fluorescence revealed by ultraviolet light exposure in Rhampholeon nalubaale sp.
nov. (UTEP 22673) from the Albertine Rift, Central Africa.

HUGHES ET AL.
476 · Zootaxa 5458 (4) © 2024 Magnolia Press
Rhampholeon bombayi sp. nov. Hughes, Dehling, Menegon, Kusamba, and Greenbaum
Bombay’s pygmy chameleon
urn:lsid:zoobank.org:act:DA3436FD-9EE8-4ED8-BC25-A1BC771C1CE8
Synonymy.
Rhampholeon boulengeri—de Witte 1965 (partim), Fischer & Hinkel 1993, Hinkel 1996, Tilbury & Tolley 2015 (partim),
Spawls et al. 2018 (partim), Tilbury 2018 (partim)
Rhampholeon sp. 3—Hughes et al. 2018
Etymology. The specific epithet honors Sidi Mubarak Bombay, an explorer of the waYao tribe who guided, led,
and interpreted for the expeditions of both Speke and Burton to discover the source of the Nile, helped Stanley find
Livingstone, and with Cameron, became the first known African to cross the African continent from east to west
(Millard 2022). Bombay was sold into slavery as a child and ended his life as the greatest African explorer of all
time.
Holotype. UTEP 21701 (field no. ELI 602), adult female, DRC, South Kivu Province, near Kalundu, 03.15552°
S, 28.42108° E, 1482 m elevation, 21 December 2010, collected by E. Greenbaum, C. Kusamba, M.M. Aristote,
and W.M. Muninga (Fig. 12A).
Paratypes. Two adult females, UTEP 21702, 22722 (field nos. ELI 617–618), DRC, South Kivu Province,
Kalundu/Mwana River, 03.15786° S, 28.4273° E, 1455 m elevation, 22 December 2010, collected by E. Greenbaum,
C. Kusamba, M.M. Aristote, and W.M. Muninga. One adult male, UTEP 22721 (field no. ELI 598), DRC, South
Kivu Province, vicinity of Kalundu, 03.15387° S, 28.42487° E, 1525 m elevation, 21 December 2010, collected by
E. Greenbaum, C. Kusamba, M.M. Aristote, and W.M. Muninga. Four adult females, UTEP 22674, 21703, 22675–
22676 (CRSN HERP 2920, 2984–2986), DRC, South Kivu Province, Mwana Kisanga, 03.15146° S, 28.44403°
E, 1529 m elevation, November 2014, collected by C. Kusamba. One adult male and three adult females, UTEP
22677–22680 (CRSN HERP 2987–2988, 2995–2996), DRC, South Kivu Province, Mwana Kisanga, 03.15678°
S, 28.43448° E, 1450 m elevation, November 2014, collected by C. Kusamba. One adult female, UTEP 22681
(CRSN HERP 2999), DRC, South Kivu Province, Hill Nkala Summit, 03.16036° S, 28.43740° E, 1555 m elevation,
November 2014, collected by C. Kusamba.
Referred specimens. UTEP 22689, 22692, 22693–22696 (field nos. JMD 2014-53, 2014-101, 2014-103–106),
RWANDA, Western Province, Nyungwe National Park, Kamiranzovu Swamp (02.46573° S, 29.15917° E, 2213
m elevation) (6 specimens). UTEP 22690–22691 (field nos. JMD 2014-55–56), RWANDA, Western Province,
Nyungwe National Park, Kamiranzovu Swamp (02.47350° S, 29.16647° E, 2330 m elevation) (2 specimens) (Fig.
12D-F). MTSN 7075–7076, RWANDA, Western Province, Nyungwe National Park (02.56753° S, 29.23079° E,
2003 m elevation) (2 specimens). MTSN 7123, RWANDA, Western Province, Cyamudongo Forest (02.54529°
S, 28.98507° E, 2033 m elevation) (1 specimen). UTEP 22697 (field no. JMD 2014-107), RWANDA, Western
Province, Cyamudongo Forest (02.53851° S, 28.99328° E, 1857 m elevation) (1 specimen). UTEP 21706, 22724–
22726 (field nos. ELI 739–741, 796), DRC, South Kivu Province, Itombwe Plateau, vicinity of Tumungu (03.53545°
S, 28.67411° E, 1835 m elevation) (4 specimens) (Fig. 12C). UTEP 21704–21705 (field nos. EBG 1286, 1346),
DRC, South Kivu Province, vicinity of Irangi (*collection notes indicate that these specimens were collected at a
nearby location by a local person [see Hughes et al. 2018]) (2 specimens) (Fig. 12B). UTEP 21708 (field no. EBG
1189), DRC, South Kivu Province, Tshibati (02.22640° S, 28.77940° E, 2030 m elevation) (1 specimen). MTSN
1640–1641 (field nos. D.C. Moyer 1640–1641), DRC, South Kivu Province, Kabobo Mountains, Kizamba River
(05.38097° S, 29.19563° E, 1904 m elevation) (2 specimens). UTEP 22682 (field no. MUSE 10154), DRC, South
Kivu Province, Itombwe Plateau, Atuyaumbu (03.56424° S, 28.25707° E, 1582 m elevation) (1 specimen). UTEP
21707, 22723 (field nos. MUSE 10146–10147), DRC, South Kivu Province, Itombwe Plateau, Mabwe (03.60583°
S, 28.34308° E, 1582 m elevation) (2 specimens).
Diagnosis. Rhampholeon bombayi sp. nov. is in the subgenus Rhinodigitum because of its distinctly bicuspid
claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily distinguishing
it from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae, R. marshalli,
R. spectrum, R. spinosus, R. temporalis, and R. viridis. Rhampholeon bombayi sp. nov. can be distinguished from
all other Rhampholeon species by the following combination of traits: (1) lack of prominent mite pockets in the
inguinal region distinguishes it from R. beraduccii, R. platyceps, R. chapmanorum, R. maspictus, R. tilburyi, R.

FIVE NEW RHAMPHOLEON FROM CENTRAL AFRICA Zootaxa 5458 (4) © 2024 Magnolia Press · 477
bruessoworum, and R. nebulauctor; (2) presence of prominent mite pockets in the axillary region distinguishes it
from R. nchisiensis and R. acuminatus; (3) distinct supra-orbital and canthal crests distinguishes it from R. hattinghi;
(4) geographic restriction to the Albertine Rift distinguishes it from R. uluguruensis, R. moyeri, R. colemani, R.
sabini, R. rubeho, R. nicolai, R. waynelotteri, and R. princeeai; (5) shorter tail length in males, genetic divergence,
and non-overlapping elevational range at parapatric sites distinguishes it from R. boulengeri; (6) larger mean body
size, longer snout length, and larger eye diameter in females, and longer inter-limb length in both sexes distinguishes
it from R. monteslunae sp. nov.; (7) slightly shorter head, snout, and mouth lengths, smaller eye diameter, shorter
inter-limb and hind limb lengths in females distinguishes it from R. nalubaale sp. nov.; (8) genetic divergence
and non-overlapping elevational range at sites of co-occurrence distinguishes it from R. plumptrei sp. nov. and R.
msitugrabensis sp. nov.
Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers
(16S, ND2, and RAG-1) among individuals of R. bombayi sp. nov. and other Rhampholeon species is presented in
Supplementary Material 1.
Description of holotype (UTEP 21701). Adult female, SVL 54.1 mm and TL 11.0 mm. Body shape leaf-like.
Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests distinct with cluster of tubercles
connected by a ridge with 13 tubercles across casque and 20 tubercles from peak-to-peak of crests. Rostral process
1.97 mm, composed of elongated tubercles. Temporal crest discrete with several enlarged tubercles extending
posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised tubercles, one raised
higher than others near snout. Ninety-eight upper and 98 lower labial tubercles present along tip of snout to rictus of
mouth. Body covered in nearly homogenous, flattened tubercles. Several larger conical tubercles present on dorsal
flanks around midbody. Crenulated dorsal crest, more prominent from mid-body to nape. Several enlarged conical
tubercles present on limbs. Claws markedly bicuspid.
Coloration of holotype (in life). A photograph of the holotype is presented in Figure 9A. Background color light
brown to gray with a darker brown hue. Limbs, tail, and top of head a darker brown color than body. Gular region
white, and this color extends from tip of chin to ventral area and towards cloaca. One prominent diagonal dark
brown line extends from near dorsal crest posteriorly to lateral flank, with a less prominent line posterior to that,
which together resemble veins on a leaf. Several small yellow tubercles present on upper lateral flanks. Largest body
tubercle near neck dark brown to black.
Variation. A summary of descriptive morphometrics for R. bombayi sp. nov. is presented in Table 3, comparative
boxplots in Figure 4, and measurements of the type specimens in Table 7. Photographs displaying color variation
in life are presented in Figure 12. Morphological proportions are generally consistent with those of the holotype.
Males have smaller body sizes (M: mean 41.5 mm, range 31.0–47.6 mm, n = 14; F: mean 47.9 mm, range 40.8–59.1
mm, n = 22) and longer tails than females (M: mean 12.9 mm, range 9.3–18.9 mm, n = 14; F: mean 11.4 mm, range
10.0–13.4 mm, n = 22). Body coloration is consistently brown to tan and/or gray with lighter orange, yellow, or red
hues. One, sometimes two or three, dark brown to red lines on the lateral flanks extend diagonally from the dorsal
crest toward the hind limbs, resembling veins of a leaf. Legs and tail coloration can be markedly darker brown than
the body. Small tubercles, especially on the eyelids, dorsal crest, and legs can often be yellowish.
Reproduction. Seven females were gravid (UTEP 21703, UTEP 22674, UTEP 22679, UTEP 22680, UTEP
22689, UTEP 22692, and UTEP 22694). Gravid females had a mean SVL of 49.1 mm (44.4–55.1 mm) and a mean
TL of 11.7 mm (10.0–13.4 mm). Most of these females were collected in November (n = 4). Two females had clutch
sizes of 4 eggs each, and one female had a clutch size of 3 eggs. The smallest specimen examined (UTEP 21708)
was collected 19 August 2007 with SVL 18.4 mm and TL 4.2 mm from Tshibati, South Kivu Province, DRC.
Distribution, natural history, and conservation. Rhampholeon bombayi sp. nov. is found in montane forests at
an elevation range of 1450–2330 m. Specimens were collected from within forests at heights of approximately 1 m
above the ground. Specimens have been collected from Kahuzi-Biega National Park, Kabobo Natural Reserve, and
Itombwe Natural Reserve, DRC, and Nyungwe Forest National Park, Rwanda. See Greenbaum & Kusamba (2012)
for detailed information on the conservation of Itombwe’s herpetofauna, and Greenbaum (2017) for a description of
the overall threats facing the herpetofauna of eastern DRC. Behavior and activity patterns are unknown, but likely
similar to that described for R. boulengeri (sensu lato) (Spawls et al. 2018; Tilbury 2018). Other lizard species
collected from montane forest in the Itombwe Plateau included Congolacerta vauereselli, Holaspis cf. guentheri,
Leptosiaphos blochmanni, L. graueri, Trachylepis varia, Trioceros johnstoni, and T. schoutedeni.

HUGHES ET AL.
478 · Zootaxa 5458 (4) © 2024 Magnolia Press
FIGURE 12. Photographs of various individuals of Rhampholeon bombayi sp. nov. in life. (A)—Adult female holotype (UTEP
21701) from Kalundu, DRC; (B)—Adult female (UTEP 21705) from the vicinity of Irangi, DRC; (C)—Adult male (UTEP
21706) from the vicinity of Tumungu, DRC; (D)—Adult female (UTEP 22690) from Nyungwe Forest National Park, Rwanda;
(E, F)—Adult male (UTEP 22691) from Nyungwe Forest National Park, Rwanda; (G)—Habitat view of the forest at the type
locality; (H)—Habitat view of the forest at Tumungu, DRC.

FIVE NEW RHAMPHOLEON FROM CENTRAL AFRICA Zootaxa 5458 (4) © 2024 Magnolia Press · 479
TABLE 7. Meristic and mensural characters in type specimens of Rhampholeon bombayi sp. nov. Linear measurements (in mm) and scale counts are given. See Materials and
Methods for explanation of character abbreviations.
UTEP
21701
Female
Holotype
UTEP
21702
Female
Paratype
UTEP
22722
Female
Paratype
UTEP
22721
Male
Paratype
UTEP
22674
Female
Paratype
UTEP
21703
Female
Paratype
UTEP
22675
Female
Paratype
UTEP
22676
Female
Paratype
UTEP
22677
Male
Paratype
UTEP
22678
Female
Paratype
UTEP
22679
Female
Paratype
UTEP
22680
Female
Paratype
SVL 54.11 42.71 40.84 44.31 52.55 48.62 47.58 43.84 42.39 47.48 55.13 50.58
TL 11.04 10.25 10.88 10.72 11.65 10.93 11.85 10.61 11.45 11.1 10.02 11.31
TaL 65.15 52.96 51.72 55.03 64.2 59.55 59.43 54.45 53.84 58.58 65.15 61.89
HL 15.51 13.06 13.02 13.52 15.1 14.36 14.27 13.82 12.7 14.19 15.32 14.47
HW 8.58 7.59 7.78 7.96 8.3 7.95 8.14 7.6 7.17 7.92 8.38 8.28
HH 7.65 6.2 6.87 7.10 7.18 7.09 7.69 7.93 6.91 7.69 7.89 7.02
ML 9.88 9.29 8.16 9.2 8.96 9.06 9.48 8.51 8.31 9.7 10.17 9.59
CE 7.06 6.41 6.39 6.4 7.11 6.63 6.89 6.52 5.82 6.48 6.8 6.57
SL 5.34 4.45 4.71 4.7 5.49 5.4 5.23 5.12 4.4 5.13 5.46 5.03
ED 4.3 3.99 3.79 4.28 4.19 4.09 4.31 4.07 3.83 4.18 4.76 4.16
CC 6.02 4.77 5.56 5.16 5.41 5.98 5.59 4.89 4.74 5.11 5.32 6.11
IL 29.52 24.06 21.51 24.11 29.25 24.69 26.93 23.6 21.22 24.79 30.12 27.27
FLL 11.05 8.83 9.7 8.89 9.93 9.5 9.6 8.98 8.4 8.94 10.01 9.66
HLL 9.67 8.05 8.5 8.68 9.39 9.16 9.05 8.35 7.12 8.75 9.44 8.4
UL 98 106 90 94 108 86 100 92 88 92 108 94
LL 98 90 86 88 102 86 98 92 84 90 94 92
RP 1.97 1.44 1.05 1.34 1.34 1.39 1.82 1.37 1.48 1.47 1.69 1.69
TL/SVL 0.204 0.239 0.266 0.242 0.222 0.225 0.249 0.242 0.270 0.234 0.182 0.224

HUGHES ET AL.
480 · Zootaxa 5458 (4) © 2024 Magnolia Press
Rhampholeon msitugrabensis sp. nov. Hughes, Kusamba, Menegon, and Greenbaum
Albertine Rift pygmy chameleon
urn:lsid:zoobank.org:act:C55B5EC4-4A4B-4D88-B318-0F122C506F33
Synonymy.
Rhampholeon boulengeri—de Witte 1965 (partim), Tilbury & Tolley 2015 (partim), Spawls et al. 2018 (partim), Tilbury 2018
(partim)
Rhampholeon sp. 4—Hughes et al. 2018
Etymology. The specific epithet is derived from the Swahili word for forest, msitu, and the German word for rift
valley, graben, in reference to the Albertine Rift forests where the species occurs.
Holotype. UTEP 21709 (field no. ELI 1159), adult female, BURUNDI, Bubanza Province, Kibira National
Park, Mpishi, 03.06177° S, 29.49343° E, 1986 m elevation, 20 December 2011, collected by E. Greenbaum, C.
Kusamba, M.M. Aristote, and W.M. Muninga (Fig. 13A).
Paratype (topotype). Same collection details as holotype, three adult females, one adult male, and one juvenile
male, UTEP 22739–22743 (field nos. ELI 1160–1164).
Referred specimen. MTSN 7213, RWANDA, Western Province, Nyungwe National Park, Mount Bigugu
(02.45565° S, 29.24927° E, 2669 m elevation) (1 specimen).
Diagnosis. Rhampholeon msitugrabensis sp. nov. is in the subgenus Rhinodigitum because of its distinctly
bicuspid claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily
distinguishing it from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae,
R. marshalli, R. spectrum, R. spinosus, R. temporalis, and R. viridis. Rhampholeon msitugrabensis sp. nov. can
be distinguished from all other Rhampholeon species by the following combination of traits: (1) lack of prominent
mite pockets in the inguinal region distinguishes it from R. beraduccii, R. platyceps, R. chapmanorum, R. maspictus,
R. tilburyi, R. bruessoworum, and R. nebulauctor; (2) presence of prominent mite pockets in the axillary region
distinguishes it from R. nchisiensis and R. acuminatus; (3) distinct supra-orbital and canthal crests distinguishes it
from R. hattinghi; (4) geographic restriction to the Albertine Rift distinguishes it from R. uluguruensis, R. moyeri,
R. colemani, R. sabini, R. rubeho, R. nicolai, R. waynelotteri, and R. princeeai; (5) slightly smaller head height
in males, genetic divergence, and allopatric distribution distinguishes it from R. boulengeri; (6) larger inter-limb
lengths in both sexes distinguishes it from R. monteslunae sp. nov.; (7) shorter inter-limb length, hind limb length,
and head length in females distinguishes it from R. nalubaale sp. nov.; (8) genetic divergence and non-overlapping
elevational range at sites of co-occurrence distinguishes it from R. plumptrei sp. nov. and R. bombayi sp. nov.
Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers (16S,
ND2, and RAG-1) among individuals of R. msitugrabensis sp. nov. and other Rhampholeon species is presented in
Supplementary Material 1.
FIGURE 13. Photograph of Rhampholeon msitugrabensis sp. nov. in life. (A)—Adult female holotype (UTEP 21709) from
Kibira National Park, Burundi.

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Description of holotype (UTEP 21709). Adult female, SVL 49.4 mm and TL 11.3 mm. Body shape leaf-like.
Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests distinct with cluster of tubercles
connected by a ridge with 10 tubercles across casque and 15 tubercles from peak-to-peak of crests. Rostral process
1.3 mm, composed of elongated tubercles. Temporal crest discrete with several enlarged tubercles extending
posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised tubercles, one raised
higher than others near snout. Ninety-six upper and 88 lower labial tubercles are present along tip of snout to rictus
of mouth. Body covered in nearly homogenous, flattened tubercles. Some larger conical tubercles present on dorsal
flanks around midbody. Crenulated dorsal crest, more prominent from midbody to nape. Enlarged conical tubercles
present on limbs. Claws markedly bicuspid.
Coloration of holotype (in life). A photograph of the holotype is presented in Figure 13A. Background color
mottled light brown to gray with several darker brown patches, especially on limbs, tail, and dorsal crest. Top of
head light brown and area near snout darker brown. Head exhibits several darker brown patches on ridges and below
eye along mouth. Gular region white, which extends from tip of chin to neck region. Two diagonal dark brown lines
extend from near dorsal crest posteriorly to lateral flanks, resembling veins of a leaf. Ventral region a much lighter
color than body.
Variation. A summary of descriptive morphometrics for R. msitugrabensis sp. nov. is presented in Table 4,
comparative boxplots in Figure 4, and measurements of the type specimens in Table 8. Morphological proportions
are generally consistent with those of the holotype. Males have smaller body sizes (M: mean 41.7 mm, range
33.6–46.8 mm, n = 3; F: mean 47.8 mm, range 45.7–49.4 mm, n = 4) and longer tails than females (M: mean 12.4
mm, range 11.5–13.7 mm, n = 3; F: mean 10.9 mm, range 10.5–11.3 mm, n = 4).
TABLE 8. Meristic and mensural characters in type specimens of Rhampholeon msitugrabensis sp. nov. Linear
measurements (in mm) and scale counts are given. See Materials and Methods for explanation of character
abbreviations.
UTEP 21709
Female
Holotype
UTEP 22739
Male
Paratopotype
UTEP 22740
Female
Paratopotype
UTEP 22741
Male
Paratopotype
UTEP 22742
Female
Paratopotype
UTEP 22743
Female
Paratopotype
SVL 49.36 44.53 49.32 33.58 45.67 46.82
TL 11.27 11.92 11.26 11.53 10.46 10.84
TaL 60.63 56.45 60.58 45.11 56.13 57.66
HL 14.54 14.42 14.38 12.07 14.07 14.01
HW 8.66 8.55 8.24 7.11 7.89 8.44
HH 7.77 8.28 7.08 6.76 7.99 6.77
ML 9.65 9.36 9.71 8.24 9.74 9.28
CE 7.18 7.27 6.86 6.09 6.97 6.69
SL 5.11 4.71 4.51 4.05 4.98 4.71
ED 4.13 4.31 4.24 3.75 3.48 4.05
CC 5.63 6.15 5.51 5.38 5.78 5.15
IL 25.79 24.79 25.74 16.79 25.5 26.15
FLL 10.13 9.97 10.14 8.03 9.8 9.79
HLL 9.2 9.37 9.82 7.02 8.79 8.91
UL 96 84 88 96 94 94
LL 88 82 86 88 88 96
RP 1.3 1.60 0.91 1.01 1.03 0.77
TL/SVL 0.228 0.268 0.228 0.343 0.229 0.232
Reproduction. Unknown. The smallest specimen examined (UTEP 22741) was collected on 20 December 2011
with SVL 33.6 mm and TL 11.5 mm from Kibira National Park, Burundi.
Distribution, natural history, and conservation. Rhampholeon msitugrabensis sp. nov. is found in montane
forests at an elevation range of 1986–2699 m. Most specimens were collected from forest edges at Mpishi near
Kibira National Park at 1986 m elevation. A single male specimen (MTSN 7213) was collected from Mt. Bigugu
in Nyungwe Forest National Park at an elevation of 2699 m. Rhampholeon bombayi sp. nov. was also found in

HUGHES ET AL.
482 · Zootaxa 5458 (4) © 2024 Magnolia Press
Nyungwe Forest National Park but allopatrically to R. msitugrabensis sp. nov. at Kamiranzovu Swamp with an
elevation range of 2003–2330 m and, also at nearby Cyamudongo Forest with an elevation range of 1857–2033
m. Specimens have been collected from two protected areas in the Rugege Highlands: Nyungwe Forest National
Park, Rwanda, and Kibira National Park, Burundi. Nyungwe Forest National Park is the largest protected area in
Rwanda, and Kibira National Park is the largest protected area in Burundi. These parks form a continuous forest
block, but the nutrient-rich soils of these highlands attract farmers in high densities, which puts pressure on the
remaining forest fragments (Burgess et al. 2007). Other lizard species collected near the type locality were typical
Albertine Rift lizard fauna, including Adolfus africanus, Chamaeleo dilepis, Congolacerta vauereselli, Hemidactylus
mabouia, Lygodactylus kibera, Kinyongia rugegensis, Trioceros ellioti, T. johnstoni, Trachylepis striata, and T.
maculilabris.
Rhampholeon monteslunae sp. nov. Hughes, Behangana, Tilbury, Dehling, Kusamba, and Greenbaum
Mountains of the Moon pygmy chameleon
urn:lsid:zoobank.org:act:9FF3B31F-E75A-414C-9986-E5D367878D43
Synonymy.
Rhampholeon boulengeri—de Witte 1965 (partim), Tilbury & Tolley 2015 (partim), Spawls et al. 2018 (partim), Tilbury 2018
(partim)
Rhampholeon sp. 5—Hughes et al. 2018
Etymology. The specific epithet is used as a noun in apposition and is derived from the Latin words for mountain,
mons, and moon, luna, in reference to the original phrase for the Rwenzori Mountains, montes lunae, or “Mountains
of the Moon”, which is the type locality for this species.
Holotype. UTEP 22735 (field no. ELI 2854), adult female, UGANDA, Western Region, Rwenzururu sub-
region, Kasese District, Rwenzori Mountains National Park, Nyakalengija entrance, 00.36029° N, 30.00922° E,
1942 m elevation, 1 June 2014, collected at night in primary forest about 1 m above the ground by D.F. Hughes, E.
Greenbaum, and M. Behangana (Fig. 14A).
Paratype (topotype). Same collection details as holotype, three adult males, one adult female, and one juvenile
female, UTEP 21389–21390, 22736–22738 (field nos. ELI 2837–2838, 2855–2857).
Paratypes. Two adult males, UTEP 21714, 22733 (field nos. ELI 2851–2852), UGANDA, Western Region,
Rwenzururu sub-region, Kasese District, Rwenzori Mountains National Park, Nyakalengija entrance, 00.36047° N,
30.00832° E, 1962 m elevation, 1 June 2014, at night in primary forest about 1 m above the ground by D.F. Hughes,
E. Greenbaum, and M. Behangana (Fig. 14C–D). One juvenile male, UTEP 22734 (field no. ELI 2853), UGANDA,
Western Region, Rwenzururu sub-region, Kasese District, Rwenzori Mountains National Park, Nyakalengija
entrance, 00.36039° N, 30.00992° E, 1936 m elevation, 1 June 2014, at night in primary forest about 1 m above the
ground by D.F. Hughes, E. Greenbaum, and M. Behangana. One adult female, UTEP 22732 (field no. ELI 2828),
UGANDA, Western Region, Rwenzururu sub-region, Kasese District, near Rwenzori Mountains National Park,
Ruboni Village, 00.34972° N, 30.02973° E, 1655 m elevation, 31 May 2014, collected at dusk about 0.5 m above
the ground on a shrub in disturbed secondary forest and garden area near Ruboni Community Hotel by D.F. Hughes,
E. Greenbaum, and M. Behangana.
Referred specimens. PEM-R 16518 (field no. CT 347), UGANDA, Western Region, Rwenzururu sub-region,
Kasese District, Rwenzori Mountains National Park, Musandama (00.73416° N, 30.17361° E, 2360 m elevation)
(1 specimen). UTEP 21710–21711, 22727 (field nos. ELI 905–907), BURUNDI, Bururi Province, Bururi Forest
Reserve (03.92857° N, 29.61662° E, 2011 m elevation) (3 specimens) (Fig. 14E–F). UTEP 22728, 21712 (field nos.
ELI 910–911), BURUNDI, Bururi Province, Bururi Forest Reserve (03.94446° N, 29.60686° E, 2201 m elevation)
(2 specimens). UTEP 22729–22731 (field nos. ELI 924–926), BURUNDI, Bururi Province, Bururi Town (03.93189°
N, 29.61846° E, 1935 m elevation) (3 specimens). UTEP 21713 (field no. CFS 1599g), BURUNDI, Bururi Province,
Bururi Forest Reserve, Rurezi River (03.93393° N, 29.61890° E, 1945 m elevation) (1 specimen).
Diagnosis. Rhampholeon monteslunae sp. nov. is in the subgenus Rhinodigitum because of its distinctly bicuspid
claws, prominent rostral process, smooth plantar surfaces, and phylogenetic placement, thus easily distinguishing
it from the six species in the other two subgenera (i.e., Rhampholeon and Bicuspis): R. gorongosae, R. marshalli,

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R. spectrum, R. spinosus, R. temporalis, and R. viridis. Rhampholeon monteslunae sp. nov. can be distinguished
from all other Rhampholeon species by the following combination of traits: (1) lack of prominent mite pockets in
the inguinal region distinguishes it from R. beraduccii, R. platyceps, R. chapmanorum, R. maspictus, R. tilburyi, R.
bruessoworum, and R. nebulauctor; (2) presence of prominent mite pockets in the axillary region distinguishes it
from R. nchisiensis and R. acuminatus; (3) distinct supra-orbital and canthal crests distinguishes it from R. hattinghi;
(4) geographic restriction to the Albertine Rift distinguishes it from R. uluguruensis, R. moyeri, R. colemani, R.
sabini, R. rubeho, R. nicolai, R. waynelotteri, and R. princeeai; (5) shorter inter-limb and snout length in males,
genetic divergence, and non-overlapping elevational range at parapatric sites distinguishes it from R. boulengeri;
(6) smaller mean body size, shorter mouth and snout lengths, smaller eye diameters, shorter inter-limb lengths, and
shorter hind limbs in females distinguishes it from R. nalubaale sp. nov.; (7) slightly shorter inter-limb lengths in
males, shorter eye diameter and inter-limb length in females, and shorter snout lengths in both sexes distinguishes it
from R. plumptrei sp. nov.; (8) smaller mean body size, shorter snout length, and smaller eye diameter in females,
and shorter inter-limb length in both sexes distinguishes it from R. bombayi sp. nov.; (9) smaller inter-limb lengths
in both sexes distinguishes it from R. msitugrabensis sp. nov.
Genetic differentiation and variation. A summary of pairwise sequence divergence for three DNA markers
(16S, ND2, and RAG-1) among individuals of R. monteslunae sp. nov. and other Rhampholeon species is presented
in Supplementary Material 1.
Description of holotype (UTEP 22735). Adult female, SVL 36.7 mm and TL 12.3 mm. Body shape leaf-like.
Casque flattened, with short head. Neck indistinct from head. Supra-orbital crests distinct with cluster of tubercles
connected by a ridge with 14 tubercles across casque and 19 tubercles from peak-to-peak of crests. Rostral process
1.34 mm, composed of elongated tubercles. Temporal crest discrete with several enlarged tubercles extending
posteriorly from mid-eye. Nares open in a posterior orientation. Canthal ridge consists of raised tubercles, one raised
higher than others near snout. Eighty-six upper and 82 lower labial tubercles present along tip of snout to rictus of
mouth. Body covered in nearly homogenous, flattened tubercles. Several larger conical tubercles present on dorsal
flanks around midbody. Weakly crenulated dorsal crest, more prominent near nape. Many enlarged conical tubercles
present on limbs. Claws markedly bicuspid.
Coloration of holotype (in life). A photograph of the holotype is presented in Figure 14A. Overall background
color tan with patches of light brown, white, and dark brown. Top of head lighter brown. Head has several darker
brown tubercles on canthal, orbital, and temporal ridges. Dark brown patch located just below eye towards edge of
mouth. Gular region white with some brown patches, which extends from tip of chin to neck region. Two diagonal
dark brown lines extend from just posterior to nape, down lateral flanks towards tail, resembling veins of a leaf. Tail
a similar brown to tan color as body. Largest body tubercles dark brown to black. Patches of orangish color present
on lateral flanks towards venter. Limbs darker brown color than body.
Variation. A summary of descriptive morphometrics for R. monteslunae sp. nov. is presented in Table 4,
comparative boxplots in Figure 4, and measurements of the type specimens in Table 9. Photographs displaying color
variation in life are presented in Figure 14. Morphological proportions are generally consistent with those of the
holotype. Males have similar body sizes (M: mean 41.1 mm, range 33.4–50.9 mm, n = 8; F: mean 40.4 mm, range
30.4–54.4 mm, n = 10), but longer tails (M: mean 13.8 mm, range 11.9–15.7 mm, n = 8; F: mean 10.9 mm, range
8.2–13.7 mm, n = 10) than females. Body coloration is consistently lighter brown to tan, usually with white hues
and two dark brown to reddish lines on the lateral flanks extending diagonally from the dorsal crest toward the hind
limbs, resembling veins on a leaf. The color of the legs can sometimes be a much darker brown than the body.
Reproduction. Unknown (but see below). The smallest specimen examined (UTEP 22736) was collected on 1
June 2014 with SVL 30.4 mm and TL 9.3 mm from Rwenzori Mountains National Park, Uganda.
Distribution, natural history, and conservation. Rhampholeon monteslunae sp. nov. is found in montane
forests at an elevation range of 1655–2360 m. Most specimens were collected from forest within Rwenzori
Mountains National Park, along the route from the Nyakalengija Gate, and a single specimen (PEM-R 16518) from
the northern end of the park near Musandama village. One specimen (UTEP 22732) was collected from secondary
forest in disturbed vegetation (ca. 0.5 m above ground) in the garden of the Ruboni Community Hotel just outside
of Rwenzori Mountains National Park. Another series of specimens were found in Bururi Forest Nature Reserve of
southern Burundi, including some from just outside the reserve around the town of Bururi. Extensive mark-recapture
chameleon surveys in October 2018 by one of us (DFH) at the type locality detected 17 individuals across seven
nights (10 males, 5 females, and 2 juveniles) that were found sleeping at a mean perch height of 627 mm (range

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FIGURE 14. Photographs of various individuals of Rhampholeon monteslunae sp. nov. in life. (A)—Adult female holotype
(UTEP 22735) from Rwenzori Mountains National Park, Uganda; (B)—Adult female (uncollected) from Rwenzori Mountains
National Park, Uganda; (C)—Adult male (UTEP 21714) from Rwenzori Mountains National Park, Uganda; (D)—Adult male
(UTEP 22733) from Rwenzori Mountains National Park, Uganda; (E)—Adult male (UTEP 21711) from Bururi Forest Reserve,
Burundi; (F)—Adult male (UTEP 21710) from Bururi Forest Reserve, Burundi; (G)—Habitat view of a riparian area near the
main trail at the type locality; (H)—Habitat view of the forest at the type locality.

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TABLE 9. Meristic and mensural characters in type specimens of Rhampholeon monteslunae sp. nov. Linear measurements (in mm) and scale counts are given. See Materials
and Methods for explanation of character abbreviations.
UTEP
22735
Female
Holotype
UTEP
22736
Female
Paratopotype
UTEP
22737
Male
Paratopotype
UTEP
22738
Male
Paratopotype
UTEP
21714
Male
Paratype
UTEP
22733
Male
Paratype
UTEP
22734
Male
Paratype
UTEP
22732
Female
Paratype
UTEP
21389*
Female
Paratopotype
UTEP
21390*
Male
Paratopotype
SVL 36.67 30.37 38.55 40.57 40.85 36.19 33.35 43.56 47 46.5
TL 12.33 9.3 12.35 12.11 13.51 13.97 11.17 11.92 12 14
TaL 50 39.67 50.89 52.68 54.36 50.16 44.72 55.48 59 60.5
HL 13.47 11.7 13.35 13.67 14.38 11.99 12.02 14.63 - -
HW 8.11 5.82 7.77 7.59 8.44 7.16 7.29 8.68 - -
HH 7.39 5.8 7.34 7.23 8.09 7.33 6.82 9.15 - -
ML 8.14 6.98 8.35 8.36 8 7.82 8.42 9.5 - -
CE 7.08 6.01 6.63 6.97 6.57 6.12 5.95 7.21 - -
SL 4.71 3.96 4.57 3.92 4.8 4.19 3.68 4.91 - -
ED 3.67 3.49 3.92 3.86 4.11 3.68 3.46 4.2 - -
CC 5.79 4.91 5.13 5.55 5.63 4.61 5.53 6.6 - -
IL 19.04 13.23 19.72 20.9 19.73 16.99 16.5 23.76 - -
FLL 10.09 7.08 8.84 9.39 9.88 8.68 8.28 11.06 - -
HLL 8.96 6.02 8.78 8.41 9.35 7.57 7.63 9.98 - -
UL 86 90 84 86 94 90 100 106 - -
LL 82 86 86 84 90 86 90 94 - -
RP 1.34 1.32 1.26 1.83 1.54 1.68 1.34 2.4 - -
TL/SVL 0.336 0.306 0.320 0.298 0.331 0.386 0.335 0.274 0.255 0.301
*Specimens measured using a standard ruler because it was used in the neuroanatomical study of Hughes et al. (2016).

HUGHES ET AL.
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254–1575 mm) and a mean perch diameter of 3.31 mm (range 1.79–4.39 mm). Two of these females were gravid,
two individuals were juveniles (SVLs 21.9 mm and 27.3 mm), and only one individual was recaptured across
seven consecutive surveys—three nights after its initial capture. This species occurs in a disjunct distribution with
relatively low intraspecific genetic diversity between allopatric populations. If populations are not discovered from
intervening sites, perhaps there were forested connections that have been lost, or fluctuations in the historical water
levels of the crater lakes led to population-level losses, which produced the disjunct distribution currently observed.
Other lizard species collected from Rwenzori Mountains National Park included Adolfus jacksoni, Kinyongia
carpenteri, K. xenorhina, K. tolleyae, Leptosiaphos meleagris, Trioceros ellioti, T. johnstoni, and T. rudis.
Discussion
The pygmy chameleon genus Rhampholeon is taxonomically difficult because all of its species are differentiated
genetically, but most are not morphologically distinct. Rhampholeon exhibit a generalized external morphology
comprising a suite of physical traits that serve to mimic a dead leaf, which is evolutionarily conserved across the
genus and several related genera (Matthee et al. 2004; Nečas & Schmidt 2004). We note that apart from a short
description of the hemipenes of R. bombayi sp. nov. from Cyamudongo Forest, Rwanda, by Tilbury (2018), there are
no descriptions of the hemipenal morphology of the other new species, and thus, additional interspecific differences
in internal anatomy for this organ may exist across the lineages. Although they are most frequently found at night
with the aid of a flashlight while sleeping in low vegetation, sometimes > 1 m off the ground, all Rhampholeon
species evolved to exploit the forest-floor niche, and thus would have experienced similarly strong selective
pressures living in the leaf litter that likely produced the extreme phenotypic conservatism of the group. In some
cases, the genetic differentiation of allopatric Rhampholeon species has not been accompanied by clear variation
in morphological traits (Branch et al. 2014; Tilbury & Tolley 2015; Menegon et al. 2022). We found that Albertine
Rift Rhampholeon represent another cryptic evolutionary radiation as revealed by the extensive overlap in our
morphological comparisons, which were based on more than 125 specimens and 13 phenotypic traits. Nevertheless,
we identified statistically significant differences in physical proportions and other morphological variation among
pygmy chameleons from this region, which can be used to distinguish the genetically differentiated populations
in combination with geography and elevation among each other and R. boulengeri. There are also differences in
climatic niche utilization among the species as indicated by their parapatric occupancy of non-overlapping elevation
zones. We note that while R. boulengeri (sensu lato) records are common in published maps from both historical
(e.g., Schmidt 1919; de Witte 1965) and contemporary sources (e.g., Spawls et al. 2018; Tilbury 2018), many of
them lack the data needed for identifications (e.g., elevation, morphology, genetics, etc.), and thus it is not possible
at present to assign them to the new species without significant uncertainty. Lastly, we documented fluorescent
tubercles of bony origin that reflect light from the ultraviolet spectrum as described by Prötzel et al. (2018), a first
for the genus Rhampholeon.
Among Albertine Rift Rhampholeon, Hughes et al. (2018) found support for a rapid radiation across the
Miocene–Pliocene boundary. Allopatric speciation via forest fragmentation during the Miocene explains the
evolution of many taxa in the region (e.g., Greenbaum et al. 2015; Hughes et al. 2017), however, this does not
fully account for the patterns within the R. boulengeri complex. The Montane Gradient Speciation Hypothesis with
parapatric speciation via niche differentiation (Moritz et al. 2000) better explains the evolution of this group because
new species seem to have formed from adaptation to different climatic regimes along an elevational gradient, where
they occupy distinct but adjacent habitats (Couvreur et al. 2021). The six species of the R. boulengeri complex, four
of which are endemic to the Albertine Rift, occur along a steep elevation gradient (500–2700 m) and exhibit genetic
differentiation. Several species in the complex occur parapatrically at non-overlapping elevational zones, particularly
at mountainous sites of co-occurrence. Pulses of forest expansion and contraction throughout the Miocene could
have produced these parapatric patterns, akin to that recently described in Afrixalus reed frogs from the Albertine
Rift (Greenbaum et al. 2022). It seems that as historical forests contracted to higher elevations during orogenic uplift
and climatic shifts, ancestral Rhampholeon populations adapted to novel physiological thresholds as they followed
the forests upwards, which ultimately would have hampered any subsequent dispersal to lower elevations during
more moist periods of high forest connectivity (Couvreur et al. 2021).

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Rhampholeon nalubaale sp. nov. exhibited a pattern of very low intraspecific genetic differentiation (16S =
0%, ND2 = 0.1–0.3%, RAG-1 = 0%, n = 10 samples) across widely separated localities (some samples are > 600 km
apart), restricted distribution in low-elevation forests, and the curious fact that only females have been found among
nearly 100 examined specimens collected from about 10 different localities. Schmidt (1919) was the first to provide
data to suggest that R. boulengeri (sensu lato) may be parthenogenetic because only females were found among the
63 specimens that he examined from two villages, both at less than 800 m elevation in the Ituri lowland rainforest
region of northeastern DRC. The results presented herein also suggest, albeit not conclusively, that R. nalubaale sp.
nov. may be parthenogenetic, providing support to the speculation of Hall (1970). Interestingly, Hall (1970) would
have had the opportunity to re-examine some material included in Schmidt (1919), but it is not clear if he did. Two
specimens currently housed at the MCZ were part of Schmidt’s (1919) collection (MCZ Herp R-13380 [AMNH
R-11592] and MCZ Herp R-13381 [AMNH R-11597]). A third specimen at the MCZ (MCZ Herp R-18375 [MCZ
FS-R6235; ZR18364]), which also could have been examined by Hall (1970), was collected in Medje, DRC, but did
not seem to come from the AMNH collection. Catalog notes by Arthur Loveridge for this specimen (“B.M. exch.”)
suggest that it came from the British Museum of Natural History, London. This specimen, nonetheless, is listed as
collected in 1914 from a site along the route of the Lang–Chapin Expedition 1909–1915, and thus was likely from
the initial collection (Schmidt 1919). We examined photographs and measurements from these three specimens at
the MCZ, which indicated that they are all female, and one (MCZ Herp R-18375) has a midventral incision, possibly
done by Hall (1970) to expose its gonads (S. Kennedy-Gold, pers. comm.).
Geographic parthenogenesis, first described by Vandel (1928), is the distinct geographic distribution of closely
related asexual lineages relative to ancestral sexual forms (Haag & Ebert 2004). Peck et al. (1998) suggested that
asexuality can be selected for in suboptimal habitats since locally adapted clones do not suffer from immigration load
in comparison to sexual genomes whose adaptation to local conditions is hampered by admixing with suboptimally
adapted immigrants from less harsh environments. This scenario could plausibly align with R. nalubaale sp. nov.
because this species occurs across a large distribution in low-elevation forests, which is distinct from its sexual
relative (R. plumptrei sp. nov.). In fact, some clades of chameleons have been found to colonize habitats distinct
from their ancestors (e.g., Bradypodion; Tolley et al. 2022a). Low-elevation sites may represent novel habitat for a
species that evolved from a high-elevation ancestor because all other lineages of the R. boulengeri complex occur
at higher elevations, thus their common ancestor may have evolved in high elevations, too (Hughes et al. 2018).
Because producing clones gives asexual species twice the reproductive potential of their sexual ancestors, this
lowland lineage may have evolved a parthenogenetic mode of reproduction from a mutation or standing genetic
variation, which could have facilitated its survival and eventual dispersal through low-elevation forests when
connections existed between mountainous regions (Glesener & Tilman 1978).
Nevertheless, other possible explanations for the pattern we detected for R. nalubaale sp. nov. described herein
include a selective sweep or a recent range expansion to account for the lack of intraspecific genetic diversity.
First, the pattern does not seem consistent with a selection event that shaped the mitochondrial genome given
that the nuclear locus also showed a similar lack of variation. Second, it seems unlikely that this lineage would
have undergone a very recent range expansion because most pygmy chameleons are diminutive, dispersal-limited
species that are usually restricted to small forest fragments (Tilbury 2018). For example, one pygmy chameleon
species was found to move an average of just 125 cm per day (range = 0–300 cm) (Foley 2002). Furthermore, it
was estimated that R. nalubaale sp. nov. diverged from its sister taxon R. plumpteri sp. nov. around 5.3 million
years ago (Hughes et al. 2018), indicating that it is a relatively ancient lineage that did not accumulate any genetic
structure during the Pleistocene as observed in many other African species (Couvreur et al. 2021). Also, very short
male life spans or sex-based niche partitioning could account for the lack of males of R. nalubaale sp. nov. found to
date. For nearly all Rhampholeon species, including the other four new species described herein and R. boulengeri,
there is apparently no pronounced niche differentiation between males and females; thus, most species seem to have
generally equal detection probabilities between sexes (Tilbury 2018). Given that our data include a negative result
(i.e., no males found to date) combined with a lack of genetic diversity across widely separated populations, it was
not possible for us to rule out other possibilities to account for the pattern observed for R. nalubaale sp. nov., such
as a bottleneck event where all loci were affected, or a novel mating tactic not previously observed among pygmy
chameleons.
Several authors have speculated that R. spectrum from West Africa is the only parthenogenetic chameleon
species (Hall 1970; Petzold 1982; Uetz et al. 2024), however, many individuals of both sexes have been found

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at numerous sites throughout its range (e.g., Wild 1994; Tilbury 2018), suggesting that this species is likely not
parthenogenetic, at least as we currently understand it. Nonetheless, given what we found, it is reasonable to think
that R. spectrum may also be a species complex as it occurs across a large geographic distribution encompassing
a wide range of elevation (Tilbury 2018), which was recently confirmed by the phylogenetic study of Tapondjou
Nkonmeneck et al. (2022). Thus, perhaps, a parallel evolutionary scenario to what we found for the R. boulengeri
complex is possible, where a cryptic pattern of speciation in R. spectrum has concealed a lineage with divergent
traits.
In addition to R. boulengeri, Steindachner (1911) described R. affinis in the same publication based on two male
specimens from Beni, North Kivu Province, DRC. This species was first synonymized with R. boulengeri by Werner
(1911) but then subsequently synonymized with the West African species R. spectrum in a checklist by Klaver &
Böhme (1997). Despite geographic and elevational overlap with the R. boulengeri complex at the type locality
(Beni, DRC [1239 m elevation]), similarities in hemipenal morphology clearly align R. affinis with R. spectrum
(Kernchen 1993). Wolfgang Böhme dissected and examined the hemipenes of one the type specimens for R. affinis
(NMW 16001: 2), during which he identified the keratinized terminal hemipenal structures typical of R. spectrum
(subgenus Rhampholeon), which are very different from the papillate horns of the R. boulengeri complex (subgenus
Rhinodigitum) (W. Bohme, pers. comm.). One of us (PW) measured the type specimens of R. affinis (NMW 16001:
1–2), and their tail lengths (19.05 mm and 21.63 mm) were longer than every specimen that we examined of the
R. boulengeri complex. Furthermore, the reported TL/SVL ratio by Tilbury (2018) for R. spectrum (up to 33%) is
much larger than that reported for the R. boulengeri complex (up to 25%), but the data sources underlying these
measurements are unclear. Additionally, our measurements of the R. affinis syntypes exhibited TL/SVL ratios of
43% and 50%, which are much more similar to the West African R. spectrum than R. boulengeri (e.g., 25–32% for R.
boulengeri syntypes [Table 2]). Taken together, it seems that R. affinis is correctly synonymized under R. spectrum,
and thus would represent the easternmost record of this species in Africa. We note that the specimens examined by
Schmidt (1919) were also collected from northeastern DRC, and thus could be affiliated with R. spectrum. However,
specimens from that collection were pictured in Schmidt (1919) and an examination of these images indicates they
are allied with a member of the R. boulengeri complex (i.e., R. nalubaale sp. nov.). For example, AMNH 11576
is pictured in life in Figure 3 of plate XXX, and AMNH 11607 is illustrated in Figures 7–8 of plate XXXII in
Schmidt (1919), and both are obviously females whose TL/SVL ratio are much more similar to R. boulengeri than
R. spectrum. In fact, the 63 specimens were measured by Schmidt (1919) and summary morphometrics are presented
on page 597 of this publication. Most mean measurements in Schmidt (1919) align with values for R. nalubaale sp.
nov. (Table 3), especially the TL/SVL ratio, for which the mean was 20% (range 17–25%) in Schmidt (1919) and
24% (range 21–27%) in our data. We also measured the TL/SVL ratios for the seven female specimens at the MCZ
examined by Hall (1970) from DRC, three of which were specimens from the initial collection of Schmidt (1919),
which resulted in a mean value of 27%—and 25% for the three Schmidt (1919) specimens only. These values show
a much greater similarity to the R. boulengeri complex than to R. spectrum (Tilbury 2018). Furthermore, Nieden
(1913), shortly after Steindachner (1911) described R. affinis, was highly doubtful that R. spectrum occurred in the
Albertine Rift region and suggested that it is likely restricted to West African forests, which raises the possibility that
the initial locality information for these specimens was incorrect. We note that, ultimately, genetic samples will be
needed to understand the species affinities and their distribution more conclusively across the region.
We wish to call attention to the threats to the biodiversity of the Albertine Rift that pose challenges to pygmy
chameleon conservation (Brooks et al. 2004; Plumptre et al. 2007; Carr et al. 2013; Ayebare et al. 2018; Greenbaum
2017; Tolley et al. 2022b). Dense human populations in the Albertine Rift have converted natural forests to agriculture
at unsustainable levels (Burgess et al. 2007; Butsic et al. 2015), a phenomenon plaguing tropical forests across
Africa (Aleman et al. 2018). Enduring civil unrest has also contributed to the degradation of the region’s natural
environments (Glew & Hudson 2007) and rendered some habitat changes irreversible (Hanson et al. 2009). Flanking
these issues are emerging threats from the extraction of recently discovered natural resources, including major
hydrocarbon projects in protected areas of the rift valley (Behangana et al. 2023). Climate change will ultimately
exacerbate these interrelated concerns by rendering natural habitats, and the species adapted to them, unsuitable
for future ecosystems (Ponce-Reyes et al. 2017). Underlying these direct threats is the rate of species discovery,
which is so slow (Fontaine et al. 2012) that many species will go extinct before they are described (Costello et al.
2013), especially in under-sampled regions such as Central Africa (Tolley et al. 2016). The description of these new
species improves our knowledge of the Albertine Rift’s true biodiversity in a manner that more optimally aligns

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conservation priorities in Africa to help regions with the greatest need. However, many of these new species have
substantially smaller geographic distributions than the area R. boulengeri (sensu lato) was previously thought to
occupy (Tilbury 2018), and thus several of them will likely be considered as threatened with extinction under the
IUCN Red List criteria (e.g., Greenbaum et al. 2022). Biodiversity discovery paired with taxonomic follow-through
in Africa, consequently, is more important for conservation now than ever before.
Acknowledgments
We first and foremost thank our Congolese and Ugandan field companions Mwenebatu M. Aristote, Wandege M.
Muninga, Jean-Pierre Mokanse, Joseph Isingoma, and Bob Katabazi. We thank Wolfgang Denzer and Wolfgang
Böhme for helpful comments, unpublished data, and fruitful discussion that improved this manuscript. We thank
Stevie Kennedy-Gold at the Museum of Comparative Zoology, Harvard University, for examining specimens,
recording measurements, and taking photographs whose inclusion improved this manuscript. The Institutional
Animal Care and Use Committee (IACUC) at UTEP approved this research (A-200902-1). Baluku Bajope of the
Centre de Recherche en Sciences Naturelles (CRSN) provided project support and permits, and the Institut Congolais
pour la Conservation de la Nature (ICCN) kindly granted permits to work in protected areas in DRC. We are
grateful for the Wildlife Conservation Society (WCS) teams who collected specimens in eastern DRC, particularly
Guillain Mitamba and Emmanuel Muhindo, and the financing by USAID, US Fish and Wildlife Service, and Arcus
Foundation that enabled the surveys to be made. We thank the Uganda National Council of Science and Technology
(UNCST) for granting us the research permit to conduct our ongoing Herpetofaunal Conservation Assessment of
Uganda (NS 481). We thank the Uganda Wildlife Authority (UWA) and the National Forest Authority (NFA) for the
necessary permits to work in the protected areas under their jurisdiction and their field staff for assisting us during
our research. We thank James Lutalo, Commissioner of Wildlife Conservation and CITES Authority for Uganda,
and Aggrey Rwetsiba, Senior Monitoring and Research Coordinator for UWA, for approval and assistance in the
export of specimens. We thank the Ngogo Chimpanzee Project for their assistance in Kibale National Park, Uganda.
Permits to enter national parks and to collect and export specimens in Rwanda were kindly issued by the Rwandan
Development Board (RDB). We thank Léonidas Nzigiyimpa of the Institut National pour l’Environnement et la
Conservation de la Nature (INECN) of Burundi for logistical support and permit negotiations. Fieldwork by DFH
was funded by the Mohamed bin Zayed Species Conservation Fund (grant 172510461) and awards provided by
UTEP, including Dodson Graduate Research Grants, Student Travel Grants, and the Dr. Keelung Hong Graduate
Research Fellowship. Fieldwork by EG in DRC was supported by the Percy Sladen Memorial Fund, an IUCN/
SSC Amphibian Specialist Group Seed Grant, K. Reed (MD), Villanova University, UTEP, National Geographic
(Research and Exploration Grant no. 8556-08), and the National Science Foundation (DEB-1145459).
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Supplementary Materials. The following supporting information can be downloaded at the DOI landing page of this
paper:
Supp. Mat. 1: Uncorrected p distances MEGA.