A new species of Chameleon (Sauria: Chamaeleonidae: Kinyongia) highlights the biological affinities between the Southern Highlands and Eastern Arc Mountains of Tanzania

Abstract

Abstract. A new species of chameleon is described from the Livingstone and Udzungwa Mountains of Tanzania. The new species is morphologically most similar to Kinyongia vanheygeni. Furthermore, a single, short rostral appendage shows the species similarity to other Eastern Arc endemic Kinyongia species (e.g. K. uthmoelleri, K. oxyrhina, K. mag-omberae and K. tenuis). Females of all these species lack any rostral ornamentation and are all very similar morpho-logically. Males of the new species, on which the morphological diagnosis is based, can be distinguished from other Kinyongia by a shorter rostral appendage that bifurcates at the tip. They are easily distinguished from K. vanheygeni, otherwise the most similar species, by differences in head scalation and the length and shape of the rostral appendage. The new species is associated with montane rainforest and is known from only four forest fragments of which two are in the Udzungwa and two in the Livingstone Mountains. Phylogenetically, the new species is sister to K. tenuis and K. magomberae, which together, form a clade that also contains K. oxyrhina. The disjunct distribution of the new species, in the Livingstone and Udzungwa mountains, stretches across the ‘Makambako Gap’ which is a putative biogeographi-cal barrier separating the distinct faunas of the Southern highlands and Eastern Arc Mountains. Evidence from this species however, points to potentially closer biological affinities between the Livingstone and Udzungwa mountains.

A new species of Chameleon (Sauria: Chamaeleonidae: Kinyongia) highlights the biological affinities between the Southern Highlands and Eastern Arc Mountains of Tanzania. Available from: https://www.researchgate.net/publication/288828844_A_new_species_of_Chameleon_Sauria_Chamaeleonidae_Kinyongia_highlights_the_biological_affinities_between_the_Southern_Highlands_and_Eastern_Arc_Mountains_of_Tanzania [accessed Jan 5, 2016].

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Acta Herpetologica 10(2): 111-120, 2015
ISSN 1827-9635 (print) © Firenze University Press
ISSN 1827-9643 (online) www.fupress.com/ah
DOI: 10.13128/Acta_Herpetol-17171
A new species of Chameleon (Sauria: Chamaeleonidae: Kinyongia)
highlights the biological anities between the Southern Highlands and
Eastern Arc Mountains of Tanzania
M M1,*, S P. L2, T R.B. D3, K M. H4, C R. T5, S
M3, K A. T5,6
1 Tropical Biodiversity Section, Museo delle Scienze, Corsodel Lavoro e della Scienza 3, 38122 Trento, Italy. *Corresponding author.
E-mail: michele.menegon@muse.it
2 Department of Life Sciences, University of Roehampton, Holybourne Avenue, Room 1053, London SW15 4JD United Kingdom
3 Wildlife Conservation Society (WCS), PO Box 1475, Mbeya & PO Box 922, Zanzibar, Tanzania
4 Department of Zoology & Wildlife Conservation, PO Box 35064, University of Dar es Salaam, Dar es Salaam, Tanzania
5 South African National Biodiversity Institute, Private Bag X7, Claremont, Cape Town, South Africa
6 Department of Botany & Zoology, University of Stellenbosch Private Bag X1, Matieland, 7602, Stellenbosch, South Africa
Submitted on 2015, 10th October; revised on 2015, 25th October; accepted on 2015, 27th October
Editor: Sebastiano Salvidio
Abstract. A new species of chameleon is described from the Livingstone and Udzungwa Mountains of Tanzania. e
new species is morphologically most similar to Kinyongia vanheygeni. Furthermore, a single, short rostral appendage
shows the species similarity to other Eastern Arc endemic Kinyongia species (e.g. K. uthmoelleri, K. oxyrhina, K. mag-
omberae and K. tenuis). Females of all these species lack any rostral ornamentation and are all very similar morpho-
logically. Males of the new species, on which the morphological diagnosis is based, can be distinguished from other
Kinyongia by a shorter rostral appendage that bifurcates at the tip. ey are easily distinguished from K. vanheygeni,
otherwise the most similar species, by dierences in head scalation and the length and shape of the rostral appendage.
e new species is associated with montane rainforest and is known from only four forest fragments of which two are
in the Udzungwa and two in the Livingstone Mountains. Phylogenetically, the new species is sister to K. tenuis and K.
magomberae, which together, form a clade that also contains K. oxyrhina. e disjunct distribution of the new species,
in the Livingstone and Udzungwa mountains, stretches across the ‘Makambako Gap’ which is a putative biogeographi-
cal barrier separating the distinct faunas of the Southern highlands and Eastern Arc Mountains. Evidence from this
species however, points to potentially closer biological anities between the Livingstone and Udzungwa mountains.
Keywords. Southern Highlands, Tanzania, Eastern Afromontane, Biodiversity, Chamaeleonidae, East Africa, new
species, reptiles.
INTRODUCTION
Exploration and subsequent research in the past
decades have substantially improved our understand-
ing of the biodiversity from the Eastern Afromontane
Region (EAR), which is known for its high species rich-
ness (Menegon and Davenport, 2008). Collectively the
Eastern Arc Mountains (EAM) and Southern Highlands
of Tanzania form a system of mountain blocks spanning
from southern Kenya, through Tanzania and into Malawi
(Lovett and Wasser, 1993). Because of the prevailing cli-
matic inuence from the warm Indian Ocean, the EAM
receives high orographic rainfall providing a relatively
stable climate. is climate stability is thought to have
reduced extinction rates for forest endemic taxa (e.g., Tol-
ley et al., 2011; Loader et al., 2014). is, coupled to ele-

112 Michele Menegon et alii
vated speciation rates as a result of specialization due to
ecotones between forest and savanna (Caro et al., 2013),
has presumably resulted in high diversity and endemism
across the EAM (Loader et al., 2015).
Our understanding of more general evolution-
ary patterns and processes for the EAM, including bio-
geographic patterns has increased dramatically for some
taxonomic groups (e.g., Tolley et al., 2011; Dimitrov et
al., 2012; Loader et al., 2014). Despite this, a number of
vertebrate species are discovered and described each year
(e.g., Rovero et al., 2014), indicating that our knowledge
is far from complete in this region. A prime example are
chameleons, for which new species are steadily being
described (Tolley and Menegon, 2013), or previously
named taxa are elevated from synonymy (Tilbury and
Emmrich 1996; Menegon et al., 2002; Mariaux and Til-
bury, 2006; Tilbury et al., 2006; Mariaux et al., 2008, Til-
bury and Tolley, 2009; Menegon et al., 2009; Greenbaum
et al., 2012; Branch et al., 2014) and these contributions
have subsequently been utilized for revealing broader
evolutionary patterns (Tolley et al., 2011; Tolley et al.,
2013; Ceccarelli et al., 2014). Essentially, the scientic
focus on EAM region has led to a substantial increase
in knowledge on the ora and fauna, but the biota of
the Southern Highlands, which is separated from the
more northern lying EAM by the dry, low-lying Makam-
bako Gap (Lovett and Wasser, 1993), is relatively poorly
known. Indeed, the Makambako gap is considered an
important turn-over region (e.g., Rovero et al., 2014) and
as a result, many biodiversity studies have instead focused
on the EAM because of its known biological wealth (e.g.
Newmark, 1998; Stanley et al., 1999).
Chameleons in the genus Kinyongia (Tilbury et al.,
2006) are a prominent group in the EAM because nd-
ings have contributed to a broader understanding of spe-
cies richness, endemism, and biogeography (e.g., Tolley et
al., 2011). Twelve of the 16 described species of Kinyon-
gia occur on isolated massifs within the EAM, with the
remainder found to the northwest in mountainous regions
of the Albertine Ri in Democratic Republic of the Con-
go, Uganda and Rwanda and on isolated volcanoes, such
as Kilimanjaro, Meru, Mt. Kenya (Tilbury, 2010; Tolley
et al., 2011; Greenbaum et al., 2012). Many species have
small distributional ranges, and are usually found on the
forested slopes of just one or a few isolated massifs. eir
isolated and restricted distributions have provided evi-
dence for a long history of persistence in EAM, and as
well as allowed inferences as to the formation and mainte-
nance of refugial areas (Tolley et al., 2011).
New biological surveys in unexplored regions such as
southern Tanzania continually reveal the presence of new
species, including Matilda’s horned viper Atheris matil-
dae (Menegon et al., 2011), the two chameleons Kinyon-
gia vanheygeni (Necas, 2009) and K. magomberae (Men-
egon et al. 2009), and Africa’s only new genus of monkey
described in the last 80 years, the kipunji Rungwecebus
kipunji (Davenport et al., 2006). Unexplored forests to the
south of the EAM (e.g. Southern Highlands) are there-
fore predicted to contain a host of species not yet known
to science. In this study, we describe a new chameleon
species in the genus Kinyongia (Fig. 1) that is found in
Afrotemperate forest from both the Livingstone (South-
ern Highlands) and Udzungwa (EAM) Mountains (Fig.
2). Using both morphological and molecular evidence,
we determine the taxonomic placement and evolution-
ary relationships for this new taxon. Both morphologi-
cal characters and genetic markers were examined and
compared to the other species of Kinyongia. Furthermore
we examine the biogeographical implications of this new
taxon given the phylogenetic hypothesis inferred from
the data.
MATERIALS AND METHODS
Material examined
e following specimens (Table 1) were examined from
the herpetological collections of the Science Museum of Trento,
Trento, Italy (MTSN and MUSE), the Department of Zoology &
Wildlife Conservation of the University of Dar es Salaam, Dar
es Salaam, Tanzania and the collection of the WCS’ Southern
Highlands Conservation Project (SHCP), Mbeya, Tanzania: K.
vanheygeni (MUSE 13523 and MUSE 13524 from Mt. Rung-
we), K. tenuis (KMH 21325 and KMH 21304 from Nilo Forest
Reserve, East Usambara Mts.), K. oxyrhina (KMH 28277 from
Ukami Forest, Udzungwa Mts.; KMH 28302 from Nyumbanitu
Forest, Udzungwa Mts., MTSN 8454 and MTSN 8412 from
Nguru South Forest Reserve), K. tavetana (MTSN 8658 and
MTSN 8661 from Kindoroko Forest Reserve, North Pare Mts.)
Molecular Analysis
To understand the phylogenetic placement the new Kinyon-
gia species a phylogenetic analysis was carried out which
included 10 individuals from the two mountain ranges, plus
multiple representatives from 17 of 19 Kinyongia species from
published datasets (Menegon et al., 2009; Tolley et al., 2011;
Greenbaum et al., 2012). e resulting dataset consisted of 47
individuals, including the outgroup taxa (Bradypodion pumi-
lum and B. melanocephalum). DNA extraction, PCR amplica-
tion, and cycle sequencing of two mitochondrial gene fragments
(ND2 and 16S) were carried out following standard proce-
dures using the following primers for ND2: L4437b and H5934
(Macey et al., 1997a, b), and 16S: L2510 and H3080 (Palumbi,
1996). Standard PCR and sequencing were followed for this
gene fragment, with PCR annealing temperature at 57˚C. All

113
A new species of chameleon from Tanzania
new sequences were deposited in European Nucleotide Archive
(Table 2).
Bayesian inference was used to investigate optimal tree
space using MrBayes 3.1.2 (Huelsenbeck and Ronquist, 2001)
for the combined mitochondrial markers (321 characters, parti-
tioned by marker: ND2, 856 bp; 16S, 465 bp), although 18 bases
were excluded for 16S due to ambiguous alignment. To inves-
tigate which evolutionary model best t the data, jModeltest
was used (Posada, 2008), and the AIC test indicated the same
model for all markers was appropriate (GTR + G). erefore,
MrBayes was run specifying six rate categories with uniform
priors for the gamma distribution for each of the partitions. To
ensure the results were robust for both datasets, the MCMC was
run twice in parallel for 10 million and 20 million generations
(four chains in each run), with trees sampled every 1000 gen-
erations. Burn-in was estimated as 1 million generations (1000
trees), as determined by examination the average standard devi-
ation of split frequencies, the convergence diagnostic (PSRF val-
ues ~ 1.0) as well as the log-probabilities and the values of each
parameter for stabilization (Ronquist and Huelsenbeck, 2003).
In addition, Tracer v1.4.1 (Rambaut and Drummond, 2007) was
used to check that the eective sample size (ESS) of all param-
eters was greater than 200 aer burn-in. A 50% majority rule
tree was constructed and nodes with ≥ 0.95 posterior probabil-
ity considered supported.
In addition to the Bayesian analysis, a maximum likeli-
hood (ML) search was run for both datasets using RAxML
HPC 7.2.8 (Stamatakis, 2006). e datasets were partitioned as
in the Bayesian analysis, with a GTR+I+G model for all mark-
ers and rapid bootstrapping halted automatically (Stamatakis et
al., 2008) is analysis was run three times to ensure that inde-
pendent ML searches produced the same topologies. We con-
sidered nodes with a bootstrap value of ≥ 70% as supported in
this analysis. Both Bayesian and likelihood analyses were run on
the CIPRES Science Gateway (Miller et al., 2010; www.phylo.
org/sub_sections/portal/). Finally, to provide a rough indica-
Fig. 1. Kinyongia msuyae sp. nov. from Livingstone Mountains in life. Pictures showing (upper) adult Male, (lower le) close up of male
head, (lower right) Adult female.

114 Michele Menegon et alii
tion of the degree of divergence between species, uncorrected
p-distances were estimated in MEGA 5.05 for 16S which had
the most complete taxon sampling (Tamura et al., 2011).
RESULTS
Molecular Analysis
The likelihood and Bayesian searches produced
the same topology and supported nodes (Fig. 3). e
phylogenetic analysis showed that the Kinyongia sam-
pled from the Livingstone and the Udzungwa moun-
tains form a well-supported clade. e uncorrected net
sequence divergence (p-distance) between this clade and
other closely related species of Kinyongia range from 1.4-
3.7% (Table 3), which is similar to within species values
of other chameleons (e.g. Menegon et al., 2009; Tilbury
and Tolley; 2009, Greenbaum et al., 2012, Tolley et al.,
2012; Branch et al., 2014). Although sequence divergence
between chameleons from Livingstone and Udzungwa
(< 1.3% for the 16S marker) are close to the lowest val-
ues found between some other Kinyongia (e.g. K. tenuis
and K. magomberae; Table 3), additional samples from
Udzungwa would be needed to obtain a more accurate
estimate of sequence divergence between the mountain
ranges. Given the genetic distinctiveness, we take the
opportunity to describe the individuals from Livingstone
and Udzungwa Mountains as a new species.
Taxonomy
Kinyongia msuyae sp. nov (Fig. 1; 3)
Holotype: Adult male in the Science Museum of
Trento, MTSN 9374 collected in Mdandu Forest Reserve,
Livingstone Mountains in January 2011 by Michele Men-
egon, Tim Davenport, Simon Loader, Sandra Dürren-
berger, Sandra Rudolf and Sophy Machaga
Type locality: Mdandu Forest Reserve, Livingstone
Mountains 1900 m above sea level, Mbeya Region, South
Eastern Tanzania (-9.769549621; 34.78832024)
K. oxyrhina
K. vanheygeni
K. magomberae
K. tenuis
K. msuyae
K. uthmoelleri
other Kinyongia species
K. cf. oxyrhina
possible dispersal routes
Makambako Gap
Udzungwa
Mts.
Southern
Highlands
(Livingstone Mts.)
Mahenge
Mts.
Kilombero
Valley
Fig. 2. Distribution of Kinyongia species in the Eastern Afromon-
tane Region. Inset map shows enlargement of Udzungwa and
Southern Highlands region of Tanzania with possible dispersal
routes of montane associated species.
Table 1. Biometrics of the holotype and paratypes. Continuous measurements given in mm.
Tot al
Length SVL TL Head
length
Head
Width
Casque
Lenght
Casque
Eye
Snouth
Lenght
Eye
Diameter
Eye-eye
Gap
Upper
Labials
Rostral
process
MTSN 9374 150.74 69.56 81.18 23.54 10.12 16.68 10.78 7.02 6.61 5.38 20 4.62
MTSN 9375 142.3 65.02 77.28 23.18 10.42 15.11 10.29 8.14 6.69 5.39 20 4.4
MTSN 9898 103.77 43.84 59.93 17.55 18.21 10.58 7.98 6.39 4.88 4.52 18 3.13
MTSN 9373 129.44 57.46 71.98 19.64 9.29 13 8.04 6.58 5.38 5.37 14 no
MTSN 9377 103.5 52.69 50.81 15.75 7.77 10.11 6.12 5.99 4.53 4.41 14 no
MTSN 7497 124.46 61.99 62.47 21.08 11.13 13.39 8.23 7.07 7.75 5.93 13 no
MTSN 9378 97.36 40.2 57.16 13.31 6.59 7.79 6.21 4.37 4.61 3.6 14 no
MUSE 13521 141. 89 68.35 73.54 22.57 11.03 14.99 10.03 7.56 7.06 5.41 19 3.63
MUSE 13522 145.53 64.25 81.28 22.37 10.31 15.66 10.36 7.14 6.39 5.01 17 4.87

115
A new species of chameleon from Tanzania
Paratype: MTSN 9375, adult male, MTSN 9373,
MTSN 9377, adult females; MTSN 9378, juvenile, same
data as holotype. MTSN 7497 collected in Sakara Nyumo
Forest Reserve (Livingstone Mountains) in January 2011
by Michele Menegon, Tim Davenport, Simon Loader and
Sophy Machaga; MTSN 8686 (adult male) collected in
Kigogo Forest Reserve, Udzungwa Mountains in Febru-
ary 2006 by Michele Menegon;
Referred material: MUSE 13521 (Field number CAM
1013) collected in Kigogo Forest Reserve by Charles A.
Msuya. MUSE 13522 (Field number KMH 28302), adult
male, collected in Nyumbanitu Forest Reserve by Louis
Hansen.
Diagnosis: A small, elongated chameleon, lacking
distinctive colours or pattern, with a tail longer than
the snout-vent length. It has a short, bone-based rostral
appendage formed by a converging, scaly elongation of
the canthi rostrales, the areas bound by the two canthi is
concave and covered in attened scales. e tips of these
elongations are free and they appear like a double-tipped
CT498
CT500
MTSN 9373
MTSN 9374
MTSN 9375
MTSN 9377
MTSN 9378
MTSN 7497
ludewa
MTSN 8686
Livingstone Mountains
Udzungwa Mountains
CAS 168917 K. tenuis
CT103 K. tenuis
MTSN 8218 K. magomberae
MTSN 8835 K. magomberae
CT192 K. oxyrhina
CT193 K. oxyrhina
CT490 K. vanheygeni
SCHP-08-R-50 K. vanheygeni
SCHP-08-R-91 K. vanheygeni
CT189 K. uluguruensis
CT191 K. uluguruensis
CAS168852 K. matschiei
CT105 K. matschiei
CAS168921 K. vosseleri
CT104 K. vosseleri
CT110 K. multituberculata
CT111 K. multituberculata
BM29
K. boehmei
JM2946 K. boehmei
CT334 K. fischeri
MTSN8490 K. fischeri
CT151 K. uthmoelleri
CT339 K. uthmoelleri
CAS201593 K. adolfifriderici
CAS201594 K. adolfifriderici
EBG2390 K. gyrolepis
EBG2391 K. gyrolepis
CT345 K. carpenteri
CT346 K. carpenteri
CT350 K. xenorhina
CT351 K. xenorhina
CT209 K. excubitor
CT106 B. melanocephalum
KT62 B. pumilum
0.05 substitutions/site
CT113 K. tavetana
CT207 K. tavetana
MTSN8661 K. tavetana
K. msuyae
Fig. 3. e best scoring maximum likelihood tree for Kinyongia, with nodes supported by maximum likelihood (bootstrap > 70%) and
Bayesian (posterior probabilities > 0.95) analyses indicated by black circles. Grey circle indicate support with Bayesian posterior probabili-
ties only.

116 Michele Menegon et alii
Table 2. Museum, GenBank and European Nucleotide Archive accession numbers (16S, ND2) for Kinyongia used in this study (CAS = Cali-
fornia Academy of Sciences; MSTN = Science Museum of Trento (formerly Museo Tridentino di Scienze Naturali); PEM = Port Elizabeth
Museum (Bayworld). N/A: sequences not available.
Species Locality ID Specimen 16S ND2
B. melanocephalum KwaZulu-Natal, South Africa CT016 N/A AY289813 HF570475
B. pumilum Western Cape, South Africa KT62 N/A AY756639 AY756689
K. adlofriderici Bwindi N.P., Uganda CAS201593 CAS201593 DQ923820 EF014304
K. adlofriderici Bwindi N.P., Uganda CAS201594 CAS201594 GQ221944 GQ221965
K. boehmei Taita Hills, Kenya BM29 N/A GQ221942 GQ221963
K. boehmei Taita Hills, Kenya JM2946 N/A GQ221948 GQ221969
K. carpenteri Rwenzori Mtns, Uganda CT345 PEM R16572 DQ923821 EF014305
K. carpenteri Rwenzori Mtns, Uganda CT346 PEM R16573 DQ923822 EF014306
K. excubitor Mount Kenya, Kenya CT209 PEM R16571 DQ923823 EF014307
K. scheri Nguru Mountains, Tanzania CT334 PEM R16566 DQ923829 EF014313
K. scheri Nguru Mountains, Tanzania MTSN 8490 MTSN 8490 GQ221951 GQ221971
K. gyrolepis Lendu Plateau, DRC UTEP 20341 UTEP20341 JN602059 JN602049
K. gyrolepis Lendu Plateau, DRC UTEP 20342 UTEP 20342 JN602055 JN602050
K. magomberae Udzungwa Mountains, Tanzania MTSN 8218 MTSN 8218 GQ221950 GQ221970
K. magomberae Magombera Forest, Tanzania MTSN 8492 MTSN 8492 GQ221952 GQ221972
K. matschiei East Usambara Mtns, Tanzania CAS 168852 CAS 168852 FR716605 FR716641
K. matschiei East Usambara Mtns, Tanzania CT105 N/A GQ221946 GQ221967
K. multituberculata West Usambara Mtns, Tanzania CT110 PEM R5735 DQ923824 EF014308
K. multituberculata West Usambara Mtns, Tanzania CT111 N/A GQ221947 GQ221968
K. msuyae Livingstone Mountains, Tanzania CT498 N/A LN997632
K. msuyae Livingstone Mountains, Tanzania CT500 N/A LN997633 LN997642
K. msuyae Livingstone Mountains, Tanzania Ludewa N/A LN997638
K. msuyae Livingstone Mountains, Tanzania MTSN7497 MTSN7497 LN997637 LN997643
K. msuyae Udzungwa Mountains, Tanzania MTSN8686 MTSN8686 LN997639
K. msuyae Livingstone Mountains, Tanzania MTSN9373 MTSN9373 LN997634 LN997644
K. msuyae Livingstone Mountains, Tanzania MTSN9374 MTSN9374 LN997635 LN997645
K. msuyae Livingstone Mountains, Tanzania MTSN9375 MTSN9375 LN997636 LN997646
K. msuyae Livingstone Mountains, Tanzania MTSN9377 MTSN9377 LN997647
K. msuyae Livingstone Mountains, Tanzania MTSN9378 MTSN9378 LN997648
K. oxyrhina Uluguru Mountains, Tanzania CT192 PEM R16569 DQ923831 EF014315
K. oxyrhina Uluguru Mountains, Tanzania CT193 PEM R16552 DQ923832 EF014316
K. tavetana Mount Kilimanjaro, Tanzania CT113 PEM R5736 DQ991233 FJ717801
K. tavetana Mount Meru, Tanzania CT207 PEM R16563 DQ923833 EF014317
K. tavetana North Pare Mountains, Tanzania MTSN 8661 MTSN 8661 FR716615 FR716649
K. tenuis East Usambara Mtns, Tanzania CAS 168917 CAS 168917 DQ923834 EF014318
K. tenuis East Usambara Mtns, Tanzania CT103 PEM R5731 DQ923835 EF014319
K. uluguruensis Uluguru Mountains, Tanzania CT189 PEM R16565 DQ923825 EF014309
K. uluguruensis Uluguru Mountains, Tanzania CT191 PEM R16557 DQ923826 EF014310
K. uthmoelleri South Pare Mtns, Tanzania CT151 PEM R16585 DQ923836 EF014320
K. uthmoelleri Mount Hanang, Tanzania CT339 N/A DQ923837 EF014321
K. vanheygeni Poroto Mountains, Tanzania CT490 LN997631 LN997649
K. vanheygeni Poroto Mountains, Tanzania SCHP-08-R-50 LN997640 LN997650
K. vanheygeni Poroto Mountains, Tanzania SCHP-08-R-91 LN997641 LN997651
K. vosseleri East Usambara Mtns, Tanzania CAS 168921 CAS 168921 GQ221943 GQ221964
K. vosseleri East Usambara Mtns, Tanzania CT104 N/A GQ221945 GQ221966
K. xenorhina Rwenzori Mtns, Uganda CT350 PEM R16570 DQ923838 EF014322
K. xenorhina Rwenzori Mtns, Uganda CT351 PEM R15568 DQ923839 EF014323

117
A new species of chameleon from Tanzania
short horn protruding over the snout by 3 to 5 mm. e
appendage is plated with subequal rounded tubercules.
Laterally, the appendage continues from the supra-orbital
crest, formed by low peaked tubercles, becoming more
serrated over the anterior rim, from where it continues
forward as a scaly rostral short horn. In the males exam-
ined it extends between 3 and 4 mm beyond the ante-
rior margin of the rostral scale. Females lack any rostral
appendage and have a lower casque.
K. msuyae does resemble K. vanheygeni Necas, 2009
and, to a lesser extent, K. uthmoelleri (Müller, 1938) in
size, general body and head shape and by possession of a
single, bone-based rostral appendage in males. It diers
from K. vanheygeni in the length of the rostral append-
age being longer, formed by more than ten scales and
pointing straight forward (less than ten scales and slight-
ly pointing upward in K. vanheygeni), from K. uthmoe-
lleri by having a horn-like longer rostral appendage (can-
thal scales in K. uthmoelleri males meet to form a ‘rostral
wall’, or protruding in form of a very short rostral pro-
jection).
Kinyongia msuyae can easily be distinguished from
the other known Kinyongia species by the combination of
the following characters: (1) presence of rostral process in
males formed by the partial fusion of the canthi rostrales
and protruding forward over the snout by 3 to 5 mm. (2)
tail longer than SVL in both sexes, and (3) gular, ventral
and dorsal crest absent.
Description of the holotype
Adult male. Total length 162.4 mm, SVL 73.7, Tail
length 88.7. Casque elongated, posteriorly raised, covered
by attened polygonal scales, giving it a smooth appear-
ance. Parietal crest formed by a series of low peaked
tuberculated scales, temporal and orbital crests present.
No occipital lobes. Nostril posteriorly directed, posi-
tioned halfway between tip of snout and the anterior rim
of the eye, and separated from upper labials by two to
three rows of attened scales. Canthi rostrales converge
above and before the nostrils in forming a single, short,
rostral appendage with two tips, giving the appearance of
two very short horns protruding beyond the rostral scale
by 4.5 mm. e rostral process is completely ossied and
covered by sub-equal, convex scales, the superior edge is
serrated. Upper labials 16, lower labials 15 on each side.
e sides of gular region is lined with 6 shallow grooves
on each side. ere is no dorsal or gular crest, while the
central part of the gular region has no groove. No signs
of dorsal, or ventral crests. Scales on body at and homo-
geneous, arranged is small clusters, those on the upper
part of the dorsum are more quadrangular and arranged
in vertical rows. Scales on limbs sub-equal, rounded, and
attened. Tail longer than the snout/vent length, later-
ally compressed, and covered by quadrangular scales
arranged in vertical rows. Hemipenes: unenverted.
Colour in preservative: e overall colour is whitish-
grey with few paler areas
Paratype variation: Paratypes show no relevant mor-
phological variation compared to the holotype. Variation
in scutellation and body proportions for the type series
and referred material are shown in Table 2.
Colour in life: K. msuyae is an overall brown to green
chameleon, sometimes with broad pale transversal bands
and scattered blue spots formed by single scales or clus-
ters of several scales. Females have oen a larger round
spot of contrasting colour on the anks (Fig. 1). e tip
of the snout, rostral appendage and limbs and top of the
casque are oen brownish to grey.
Distribution: Refer to Figure 2.
Etymology: e species is named aer and dedicated
to Charles A. Msuya, a pioneer of Tanzanian herpetology,
who collected the rst known specimen attributable to
this species and has spent most of his life studying Tan-
zanian wildlife.
DISCUSSION
The phylogenetic analyses, sequence divergence
estimates, and morphological assessment suggest that
there is a previously unknown but distinctive species of
Kinyongia from the Livingstone and Udzungwa moun-
tains, which we describe as Kinyongia msuyae. is spe-
cies is sister to chameleons found in the Eastern Arc
Mountains (i.e. K. tenuis from Usambara Mountains, K.
magomberae from Udzungwa Mountains, and K. oxy-
rhina from Uluguru Mountains). There was no clear
morphological differentiation between the population
on the Udzungwa and that from Livingstone Mountains,
despite these two mountain ranges being separated by
Table 3. Sequence divergence (p-distances) for Kinyongia within
species (on diagonal) and between selected species for 16S (lower
matrix). Populations of K. msuyae from Livingstone and Udzungwa
are given separately. N/A = not available.
12345
1K. msuyae (Livingstone) 0.0006    
2K. msuyae (Udzungwa) 0.0129 N/A   
3K. tenuis 0.0262 0.0292 0.0000  
4K. oxyrhina 0.0208 0.0235 0.0246 0.0000 
5K. magomberae 0.0269 0.0370 0.0137 0.0186 0.0000

118 Michele Menegon et alii
the Makambako Gap (ca. 150 km apart). Sequence diver-
gence between chameleons from these localities less than
what is normally found between species. Population level
dierences may exist, but additional sampling would be
required to conrm that hypothesis.
e close relationship among populations has some
important biogeographic implications. The Southern
Highlands have long been regarded as isolated and not
part of the Eastern Arc Mountains, with the Makambako
gap considered inhospitable, preventing dispersal. How-
ever, recent molecular data has started to alter this view.
Phylogenetic analyses for the shrew, Myosorex, suggest
that the Makambako Gap is of little consequence in the
historical biogeography of the genus (Stanley and Ess-
eltyn, 2010). Similarly, there is little morphological vari-
ation among populations of the murid rodent Hylomys-
cus arcimontensis on either side of the Makambako Gap
(Carleton and Stanley, 2005), and the newly discovered
kipunji monkey (Rungwecebus kipunji) has populations
on both sides of the gap (Jones et al., 2005; Davenport et
al., 2006). ere are also some commonalities in the avi-
fauna among populations on the Nyika Plateau, Mount
Rungwe, and the southern Udzungwas, with no evidence
of the Makambako Gap having a biogeographic inuence
(Stuart et al., 1993). Furthermore the Southern Highlands
might have served as a dispersal route for amphibians,
connecting the Udzungwa and the Mahenge Mts, the two
southernmost mountain blocks of the Eastern Arc (Men-
egon et al., 2011; Loader et al., 2014).
Interestingly, from a biogeographic perspective, the
most suitable dispersal route for forest endemics from
Udzungwa and Mahenge mountains – both part of the
Eastern Arc Mountains - does not appear to be the short-
est straight line distance, which would require crossing
the Kilombero Valley (an ancient, deep, wide valley).
Instead, the continuous ridge of highlands connecting the
southern Udzungwa, through the Southern Highlands/
Livingstone Mountains via the Makambako Gap and then
northeast to the Mahenge Mountains may have remained
more suitable over historical times, potentially with for-
ested areas (see Fig. 2).
e description of Kinyongia msuyae provides a tan-
talizing piece of evidence suggesting strong biogeographi-
cal anities between the Southern Highlands (i.e. Liv-
ingstone Mountains) and the Eastern Arc Mountains (i.e.
Udzungwa Mountains). e Makambako Gap may not be
a turn-over region of high signicance between the EAM
and the Southern Highlands, rejecting previous claims
of its biogeographical importance as a barrier. Instead,
it is likely that some taxa can or have crossed this bar-
rier, or that the gap was formerly less dry, forming a cor-
ridor between Udzungwa and the Southern Highlands.
Our increased understanding of the Southern Highlands
is revealing that the region is more species rich than had
been supposed, possibly similar in scale to some of the
Eastern Arc Mountain forests.
ACKNOWLEDGEMENTS
For advice, help with eldwork, granting national
and local permits for research and export in Tanzania,
we thank (no particular order) Tanzania Commission
for Science and Technology (COSTECH research per-
mit RCA 2001-272; RCA 2007-153, RCA 2009-306-NA-
2009-201, 2011-239-NA-2011-82, 2006 and 2007-72-Na-
2006-19), Tanzania Wildlife Research Institute (TAWIRI),
Wildlife Division.
We are also grateful to many people and organiza-
tions that provided assistance in the eld, logistical sup-
port and advice, including Sandra Dürrenberger, San-
dra Rudolf, Noah Mpunga, Tanzania Forest Conserva-
tion Group, and colleagues of the Wildlife Conservation
Societies Southern Highlands Conservation Programme.
anks to Nicholas Barbieri for the help in the lab. is
work was supported by the Swiss National Science Foun-
dation (grant number 31003A-133067 to SPL), Swiss
Academy of Sciences, Freiwillige Akademische Gesells-
cha Basel, e University of Basel, and the South Afri-
can National Biodiversity Institute. MM is grateful to the
Gino Zobele Fund for Research and the Lipparini family
for their generous support. Phylogenetic analyses were
run at the Cyberinfrastructure for Phylogenetic Research
Science Gateway v 3.3 (CIPRES).
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