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


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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: [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)
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.
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.
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.
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
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).
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.
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
(Livingstone Mts.)
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
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
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
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
MTSN 9373
MTSN 9374
MTSN 9375
MTSN 9377
MTSN 9378
MTSN 7497
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
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
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-
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.
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.
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.
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
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... Various searches in isolated forest patches of East and Central Africa have revealed distinct species of Kinyongia (e.g. Lutzmann & Nečas, 2002;Menegon et al., 2009Menegon et al., , 2015Nečas, 2009;Nečas et al., 2009), including two new species from the AR (Greenbaum et al., 2012a;. Therefore, we anticipate that additional undescribed Kinyongia lineages occur in other poorly explored forest fragments of the AR. ...
... Differential diagnosis: A medium-sized forest chameleon that is distinguished from most congeners by the absence of a rostral process in both sexes [K. asheorum, K. boehmei (Lutzmann & Nečas, 2002), K. carpenteri, K. fischeri, K. magomberae Menegon et al. (2009), K. matschiei (Werner, 1895), K. msuyae Menegon et al. (2015), K. multituberculata (Nieden, 1913), K. oxyrhina (Klaver & Böhme, 1988), K. tavetana (Steindachner, 1891), K. tenuis (Matschie, 1892), K. uluguruensis (Loveridge, 1957), K. uthmoelleri (Müller, 1938), K. vanheygeni Nečas, 2009, K. vosseleri (Nieden, 1913 ...
... Results presented here are generally consistent with the phylogenetic relationships within Kinyongia recovered by Mariaux, Lutzmann & Stipala (2008), Menegon et al. (2009), Greenbaum et al. (2012a, Tolley et al. (2013), Menegon et al. (2015) and . Also, these findings support the phylogenetic positions of K. msuyae and K. mulyai, two recently described forest chameleon species . ...
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The Albertine Rift (AR) is a centre for vertebrate endemism in Central Africa, yet the mechanisms underlying line-age diversification of the region's fauna remain unresolved. We generated a multilocus molecular phylogeny consisting of two mitochondrial (16S and ND2) and one nuclear (RAG1) gene to reconstruct relationships and examine spatiotemporal diversification patterns in the AR endemic forest chameleon, Kinyongia adolfifriderici (Sternfeld, 1912). This widely distributed species was revealed to be a complex of four genetically distinct and geographically isolated species. Three new species are described based on molecular analyses and morphological examinations. We find that K. rugegensis sp. nov. (Rugege Highlands) and K. tolleyae sp. nov. (Kigezi Highlands) form a well-supported clade, which is sister to K. gyrolepis (Lendu Plateau). Kinyongia itombwensis sp. nov. (Itombwe Plateau) was recovered as sister to K. adolfifriderici (Ituri Rainforest). The phylogeographic patterns we recovered for Kinyongia suggest that speciation stemmed from isolation in forest refugia. Our estimated diversification dates in the Miocene indicate that most species of Kinyongia diverged prior to the aridification of Africa following climate fluctuations during the Pleistocene. Our results highlight the AR as a focal point of diversification for Kinyongia, further elevating the global conservation importance of this region.
... Menegon et al., 2011;Rovero et al., 2014). The rate of new reptile descriptions in Africa shows little indication of reaching a plateau (Menegon et al., 2015), and species numbers have increased by 65% in the last 26 years (Branch unpubl. obs.). ...
... Broadley, 2006;Bauer and Menegon, 2006) indicating that they contain hidden diversity. For instance, the biodiversity wealth of Eastern Arc Mountains is well known due to the extensive scientific focus it has obtained, but the Southern Highlands, to the south of Eastern Arc Mountains, divided by the Makambako gap, remains poorly known and has stronger affinities to the Eastern Arc than was previously acknowledged (Menegon et al., 2015). ...
... Systematic studies of East African members of the genus Rhampholeon [1,6] and other chameleons [70,71] have resulted in recently described species, all of which relied, in part, on phylogenetic support and levels of genetic divergence (uncorrected p-distances), similar to those reported here in Table 1, as justification for their formal taxonomic recognition. Based on the geographic distribution of genetic diversity we recovered, our results support a range of possibilities, likely for at least two potentially cryptic species within the R. spectrum complex. ...
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Pygmy Chameleons of the genus Rhampholeon represent a moderately diverse, geographically circumscribed radiation, with most species (18 out of 19 extant taxa) limited to East Africa. The one exception is Rhampholeon spectrum, a species restricted to West-Central African rainforests. We set out to characterize the geographic basis of genetic variation in this disjunctly distributed Rhampholeon species using a combination of multilocus Sanger data and genomic sequences to explore population structure and range-wide phylogeographic patterns. We also employed demographic analyses and niche modeling to distinguish between alternate explanations to contextualize the impact of past geological and climatic events on the present-day distribution of intraspecific genetic variation. Phylogenetic analyses suggest that R. spectrum is a complex of five geographically delimited populations grouped into two major clades (montane vs. lowland). We found pronounced population structure suggesting that divergence and, potentially, speciation began between the late Miocene and the Pleistocene. Sea level changes during the Pleistocene climatic oscillations resulted in allopatric divergence associated with dispersal over an ocean channel barrier and colonization of Bioko Island. Demographic inferences and range stability mapping each support diversification models with secondary contact due to population contraction in lowland and montane refugia during the interglacial period. Allopatric divergence, congruent with isolation caused by geologic uplift of the East African rift system, the "descent into the Icehouse," and aridification of sub-Saharan Africa during the Eocene-Oligocene are identified as the key events explaining the population divergence between R. spectrum and its closely related sister clade from the Eastern Arc Mountains. Our results unveil cryptic genetic diversity in R. spectrum, suggesting the possibility of a species complex distributed across the Lower Guinean Forest and the Island of Bioko. We highlight the major element of species diversification that modelled today's diversity and distributions in most West-Central African vertebrates.
... Mulanje) and the Eastern Arc Mountains of Tanzania (Lawson 2013). Several biogeographic connections have also been found across one of the main biogeographic breaks separating the Eastern Arc Mountains from the Southern Highlands (i.e., the Makambako Gap) in mammals (e.g., Lophocebus; Palmer 1903, Jones et al. 2005), chameleons (e.g., Kinyongia;Tillbury et al. 2006, Menegon et al. 2015, and vipers (e.g., Atheris; Cope 1862, Menegon et al. 2011). Different clustering methods suggest different ways of subsetting the Eastern Arc, and a combination of these methods confirms the presence of multiple distinct subregions and transitional flanking regions that parallels other biogeographic assessments within the region (Dowsett 1986, Cordeiro 1998. ...
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The Afromontane mountains are a complex series of highlands that have intermittently been connected by habitat corridors during climatic cycles, resulting in a mosaic of range disjunctions and allospecies complexes in the present day. Patterns of community relatedness between geographic regions are often determined through single-species analyses or spatial analyses of diversity and nestedness at the species level. To understand patterns of Afromontane community evolution and to assess the effects of taxonomy on our understanding of biogeographic patterns, I concatenated three lists of Afromontane bird taxa divided into five taxonomic hierarchies. These lists were converted into a presence-absence matrix across 42 montane regions and analyzed using a variety of clustering techniques based on a replicable coding pipeline. I used these lists and methods to determine patterns of relatedness between montane blocks, to assess the consistency with which biogeographic regions were recovered, and to shed light on the patterns of connectivity within the Afromontane region. My results reaffirm the distinctiveness of many biogeographic regions (e.g., the Cameroon Highlands) while also clarifying regional relationships and the presence of 'transition zones' between regions. Differences between lists illustrated how our understanding of taxonomy and distribution in the Afromontane highlands can also change our understanding of Afromontane biogeography. Most notably, I found evidence for an Expanded Eastern Arc that included the Eastern Arc Mountains and highlands in Malawi, Mozambique, and Zimbabwe. This study presents a rigorous yet easily adjustable pipeline for studying regional biogeography from multiple taxonomic perspectives using both traditional and novel approaches.
... There is no increase in tracheal diameter and the hyaline cartilage rings are equally spaced. This is known to represent the condition for the vast majority of the 213 currently recognized chameleon species (Glaw, 2015;Hughes, Kusamba, Behangana, & Greenbaum, 2017;Menegon et al., 2015;Prötzel, Lambert, et al., 2018;Prötzel, Vences, et al., 2018; Figure 1c shows the gross anatomy of the throat region of a male veiled chameleon, C. calyptratus. The three abovementioned muscles are present, but here each directly contacts the gular pouch, as does the posterior musculature of the tongue, comprised of the M. accelerator linguae and the M. hyoglossus. ...
Numerous chameleon species possess an out‐pocketing of the trachea known as the gular pouch. After surveying more than 250 specimens, representing nine genera and 44 species, we describe two different morphs of the gular pouch. Species of the genera Bradypodion and Chamaeleo, as well as Trioceros goetzei, all possess a single gular pouch (morph one) formed from ventral expansion of soft tissue where the larynx and trachea meet. Furcifer oustaleti and F. verrucosus possess from one to four gular pouches (morph two) formed by the expansion of soft tissue between sequential hyaline cartilage rings of the trachea. In T. melleri, examples of both morphs of the gular pouch were observed. Morphometric data are presented for 100 animals representing eight species previously known to possess a gular pouch and two additional species, B. thamnobates and B. transvaalense. In the species with the absolutely and relatively largest gular pouch, C. calyptratus, a significant difference was found between sexes in its width and volume, but not its length. In C. calyptratus, we show that an inflated gular pouch is in contact with numerous hyoid muscles and the tongue. Coupled with the knowledge that C. calyptratus generates vibrations from the throat region, we posit that the tongue (M. accelerator linguae and M. hyoglossus) and supporting hyoid muscles (i.e., Mm. sternohyoideus profundus et superficialis and M. mandibulohyoideus) are involved in the production of vibrations to produce biotremors that are amplified by the inflated gular pouch and used in substrate‐borne communication.
... Of the 210 species of chameleons described so far, 90 species are endemic to Madagascar (Glaw 2015;Menegon et al. 2015;Hughes et al. 2017;Prötzel et al. 2017Prötzel et al. , 2018. Although there is already a great chameleon diversity on the island, new species are being discovered regularly (e.g. ...
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The taxonomy of the Malagasy chameleon Furcifer rhinoceratus (Gray, 1845) is poorly resolved. The aim of this study is to clarify the taxonomic status of Chamaeleon voeltzkowi Boettger, 1893 and Chamaeleon monoceras Boettger, 1913 both only known from single or very few specimens mostly collected more than 100 years ago and currently considered as synonyms of Furcifer rhinoceratus. Using osteological data from micro-X-ray computed tomography (micro-CT) combined with traditional morphological characters and morphometrics we resurrect both taxa from the synonymy of F. rhinoceratus as F. voeltzkowi and F. monoceras, respectively. Compared to F. rhinoceratus, F. monoceras is smaller, has a relatively shorter tail, a longer and thinner rostral appendage, a poorly developed gular crest and no ventral crest, whereas F. voeltzkowi has a smaller rostral appendage, higher casque and the dorsal crest is continuous with the tail crest. Compared to the broad rostral appendage formed by the anterior protuberance of the premaxillary process of the maxilla, which has serrated edges in F. rhinoceratus, F. monoceras presents a long rostral appendage with a smooth dorsal edge that progressively narrows, and the nasal aperture is extended along the elongated appendage; F. voeltzkowi presents a smaller but curved rostral appendage with a crenate edge. The prefrontal and postorbitofrontal approach one another forming a large, laterally closed supraorbital fontanelle in F. rhinoceratus while in F. monoceras they do not approach, leaving a laterally open fontanelle, and in F. voeltzkowi the fontanelle is diminutive. Furcifer voeltzkowi also differs from the similar F. labordi by a smaller size of the rostral appendage, less bulging casque and body pholidosis. The former exhibits a conspicuous white lateral band comprising heterogeneous scalation. Furcifer labordi, on the other hand, has a homogeneous scalation with a remarkable reticulate pattern. Osteologically, the shape of the prefrontal and the connection of the postorbitofrontal with the parietal also differ greatly between the two. Using micro-CT scans we detected key differences that would be otherwise impossible to determine. We also provide a brief morphological and osteological description of the species and strongly recommend efforts to rediscover these two poorly known taxa in order to enable additional studies and to assess their conservation status.
... The precise number of reptile species in the EACF is unknown, as new species continue to be described (e.g. Bauer & Menegon, 2006;Mariaux & Tilbury, 2006;Menegon et al., 2009Menegon et al., , 2011bMenegon et al., , 2015Fisseha et al., 2013;Malonza & Bauer, 2014). Myers et al. (2000) reported 188 reptile species in the EACF, 50 of which were endemic. ...
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We present an account of the 909 globally threatened taxa (793 species, 74 subspecies, 42 varieties) of animals and plants in the Eastern Arc Mountains and Coastal Forests of Kenya and Tanzania and the sites in which they occur based upon a review of the 2015 IUCN Red List of Threatened Species. Results for animals are summarised by Class (Amphibia, Aves, Gastropoda, Insecta, Malacostraca, Mammalia, Reptilia) and presented for plants as a whole (Classes Bryopsida, Cycadopsida, Jungermanniopsida, Liliopsida, Lycopodiopsida, Magnoliopsida, Pinopsida, Polypodiopsida). We analyse the status of previously known and newly identified sites in which globally threatened biodiversity occurs and summarise the current state of research on the globally threatened and ecologically critical biodiversity of the EACF. We then provide recommendations for future research, environmental regulations, and management regimes based upon comprehensive and reliable data to ensure the continued survival of the EACF's biodi...
Whether the Colobus angolensis that reside in the fragmented forests in eastern Kenya and Tanzania represent one subspecies or two has been debated for 50 years. Morphological and more recent genetic and ecological studies suggest that these populations represent two subspecies, C. a. palliatus and C. a. sharpei. However, their distribution of mitochondrial variation remains unresolved since the genetic study only characterized four populations at the range ends. Therefore, we characterized five populations in the area of the hypothesized subspecies divide. We identified eight new haplotypes which, combined with those previously identified, provided 26 haplotypes from nine populations for analysis. Haplotypes found south of the Rufiji River cluster together but separately from northern haplotypes. The largest sequence differences within cytochrome b occur between population pairs representing opposite sides of the river; their mean difference (1.5%) is more than that of other primate subspecies. Analysis of molecular variance attributes most of the variation to that north versus south of the river. These results support the previous subspecies distinction between C. a. palliatus (northern) and C. a. sharpei (southern), divided by the Rufiji River. The estimated time of the most recent common ancestor of all haplotypes indicates that the subspecies have been isolated from each other for approximately 550,000 years. The common ancestor of northern and southern haplogroups was 370,000 and 290,000 years ago, respectively. Nevertheless, the correlation between genetic and geographic distances suggests that isolation‐by‐distance contributed to population structuring. Significant variation among populations, with only three haplotypes shared between populations, also indicates that an extended period of isolation drove population distinctiveness. Considering these results, we evaluate hypotheses about the founding and differentiation of these subspecies during Pleistocene climatic fluctuations and propose a novel, more direct migration route from Central Africa to their current range navigating Lake Tanganyika, the central Tanzanian corridor, and the Rufiji River. Proposed diversification of Colobus angolensis in the eastern forests of Kenyan and Tanzania. The ancestor to these colobus populations arrived in Tanzania sometime before 550,000 years ago. They began diverging into two different subspecies 290,000–550,000 years with the Rufiji River as the putative boundary. Since then, the populations north of the river and the populations south of the river have continued to diverge due to climatic changes during the Pleistocene resulting in a highly fragmented landscape in and around the Eastern Arc Mountains. Results support two colobus subspecies (Colobus angolensis palliatus and C. a. sharpei) in the eastern forests of Kenya and Tanzania, having diverged approximately 550,000 years ago, with the Rufiji River as the putative boundary that divides them. Each population has predominantly unique mitochondrial haplotypes, indicating the importance of protecting these populations to maintain subspecies diversity. Based on their phylogeographic patterns, a novel migration route for colobus is proposed from the Congo Basin to their current range navigating Lake Tanganyika, the central Tanzanian corridor, and the Rufiji River. Results support two colobus subspecies (Colobus angolensis palliatus and C. a. sharpei) in the eastern forests of Kenya and Tanzania, having diverged approximately 550,000 years ago, with the Rufiji River as the putative boundary that divides them. Each population has predominantly unique mitochondrial haplotypes, indicating the importance of protecting these populations to maintain subspecies diversity. Based on their phylogeographic patterns, a novel migration route for colobus is proposed from the Congo Basin to their current range navigating Lake Tanganyika, the central Tanzanian corridor, and the Rufiji River.
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The Livingstone Mountains (LM; also known as the Kipengere Range) found in southwestern Tanzania at the northern end of Lake Nyasa are an important region for understanding the biogeography of Eastern Africa. The two branches of the East African Rift Valley meet here and the mountains might represent stepping stones for colonization and migration between different parts of the Eastern Afromontane Biodiversity Hotspot (especially the link between the Eastern Arc Mountains, EAM, and the Southern Rift Mountains, SRM), as well as an efficient barrier to gene flow for taxa living in drier savannahs in lower elevations. Here we combine new mitochondrial sequence data from 610 recently sampled rodents and shrews with available georeferenced genetic data (3538 specimens) from southern Tanzania, northern Malawi/Zambia and northern Mozambique and compare the spatial genetic structure among different taxa. There is no universal phylogeographic pattern in taxa preferring humid montane habitats. For some of them, the Makambako Gap acts as a barrier between the SRM and the EAM, but other taxa can bridge this gap. Barriers within the EAM (frequently) and within the SRM (sometimes) appear more important. The Rukwa rift between the SRM and the ARM is an important barrier that perhaps can only be crossed by taxa that are not that strictly tied to humid montane environments. For mammals living in lower-elevation savannah-like habitats, the LM can act as a strict barrier to gene flow, and together with the Ufipa Plateau, Lake Nyasa and the EAM create a very similar phylogeographic pattern with three recognizable genetic groups in most savannah-dwellers. The Livingstone Mountains thus appear to be one of the most important biogeographic crossroads in Eastern Africa.
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A new species of chameleon, Trioceros wolfgangboehmei sp. nov. , inhabiting the northern slopes of the Bale Mountains in Ethiopia, is described. It differs from its Ethiopian congeners by a combination of the following features: presence of a prominent dorsal crest with a low number of enlarged conical scales reaching along the anterior half of the tail as a prominent tail crest, a casque raised above the dorsal crest, heterogeneous body scalation, long canthus parietalis, rugose head scalation, high number of flank scales at midbody and unique hemipenial morphology. Based on morphological characteristics, phylogenetic discordances of previous studies and biogeographical patterns, this new species is assigned to the Trioceros affinis (Rüppell, 1845) species complex. An updated comprehensive key to the Trioceros found in Ethiopia is provided.
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The genus Chamaeleo, currently subdivided into two sub-genera, Chamaeleo (Chamaeleo) and Chamaeleo (Trioceros) (Klaver & Böhme 1986), is reviewed from both a morphological and genetic basis and it is concluded that the two subgenera are sufficiently distinct as to warrant their formal elevation to seperate and distinct genera. Evaluation of the soft anatomy and several other characters provide sufficient basis for making this distinction. The proposed change is supported by the demonstration of monophyletic groupings (based on two mitochondrial and one nuclear gene) consistent with distinct genera.
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Background The East African spiny-throated reed frog complex (Hyperolius spinigularis, H. tanneri, and H. minutissimus) is comprised of morphologically similar species with highly fragmented populations across the Eastern Afromontane Region. Recent genetic evidence has supported the distinctiveness of populations suggesting a number of cryptic species. We analyse newly collected morphological data and evaluate the taxonomic distinctiveness of populations. Results We find three new distinct species on the basis of morphological and molecular evidence. The primary morphological traits distinguishing species within the Hyperolius spinigularis complex include the proportions and degree of spinosity of the gular flap in males and snout-urostyle length in females. Other features allow the three species to be distinguished from each other (genetics). We refine the understanding of H. minutissimus which can be found in both forest and grassland habitats of the Udzungwa Mountains, and provide more details on the call of this species. Further details on ecology are noted for all species where known. Conclusions Three new species are described and we narrow the definition and distribution of Hyperolius spinigularis and H. minutissimus in East Africa. The spiny-throated reed frogs have highly restricted distributions across the fragmented mountains of the Eastern Afromontane region. Given the newly defined and substantially narrower distributions of these spiny-throated reed frog species, conservation concerns are outlined. Electronic supplementary material The online version of this article (doi:10.1186/s13104-015-1050-y) contains supplementary material, which is available to authorized users.
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Detailed knowledge of species distributions, endemism patterns and threats is critical to site prioritization and conservation planning. However, data from biodiversity inventories are still limited, especially for tropical forests, and even well recognized hotspots remain understudied. We provide an example of how updated knowledge on species occurrence from strategically directed biodiversity surveys can change knowledge on perceived biodiversity importance, and facilitate understanding diversity patterns and the delivery of conservation recommendations.
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The taxonomic status of recently discovered populations of pygmy chameleons (Rhampholeon) from the northern Mozambique montane isolates of Mt. Chiperone, Mt. Mabu, Mt. Inago and Mt. Namuli are assessed, and compared with the closest geographical congeners, including Rhampholeon platyceps Günther 1893 from Mt. Mulanje, and Rh. chapmanorum Tilbury 1992 from the Malawi Hills, both in southern Malawi. Relationships were examined using morphological features and a phylogenetic analysis incorporating two mitochondrial and one nuclear marker. The phylogeny showed that each montane isolate contained a distinct, well-supported clade of chameleons. Chameleons from the Mozambican montane isolates are within a monophyletic clade inclusive of species from southern Malawi (Rh. platyceps and Rh. chapmanorum). Although some relationships are unresolved, the southern Malawi and Mozambican isolates appear to share their most recent common ancestor with species from the Eastern Arc Mountains and Southern Highlands of Tanzania and Malawi (Rh. moyeri, Rh. uluguruesis, Rh. nchisiensis). Along with Rh. beraduccii and Rh. acuminatus, all are included in the subgenus Rhinodigitum. Sister to this larger clade are species from west/central Africa (Rh. temporalis, Rh. spectrum) and the Rh. marshalli-gorongosae complex from southwest Mozambique and adjacent Zimbabwe. Morphological and molecular results confirm that Brookesia platyceps carri Loveridge 1953 is a junior subjective synonym of Rhampholeon platyceps Günther 1892. Historical records of Rh. platyceps from the Shire Highlands (Chiromo) and the Zomba Plateau, are incorrect and the species is now considered endemic to the Mulanje massif. All of the four newly discovered, isolated populations are genetically and morphologically distinct, and we take the opportunity to describe each as a new species. Rhampholeon (Rhinodigitum) maspictus sp. nov. is restricted to Mt. Mabu and distinguished by its large size, well-developed dorsal crenulations, and bright male breeding coloration; Rhampholeon (Rhinodigitum) nebulauctor sp. nov. is restricted to Mt. Chiperone and distinguished by its small size, weakly-developed dorsal crenulations, and a large rostral process in males; Rhampholeon (Rhinodigitum) tilburyi sp. nov. is restricted to Mt. Namuli and distinguished by its small size, weakly-developed dorsal crenulations, and prominent flexure of the snout in males; and Rhampholeon (Rhinodigitum) bruessoworum sp. nov. is restricted to Mt. Inago and distinguished by its small size, weakly-developed dorsal crenulations, large rostral process in males, and relatively long tail in both sexes.
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AimThe persistence and stability of habitats through time are considered predictors of high levels of biodiversity in some environments. Long-term habitat persistence and stability may explain the species-rich, endemic forest fauna and flora of the Eastern Afromontane Biodiversity Region (EABR). Using complementary phylogenetic and biogeographical approaches, we examine evolutionary patterns in EABR brevicipitid frogs. Using these data, we test whether brevicipitid history reflects patterns of long-term forest persistence and/or stability across the EABR.LocationEast Africa.MethodsA dated phylogeny for brevicipitids was constructed using two nuclear and three mitochondrial markers. Alternative diversification models were used to determine signal for constant or varying net diversification rates. Using our dated tree, we identified areas of high phylogenetic diversity (PD), and inferred ancestral areas using likelihood and Bayesian approaches.ResultsBrevicipitids have a long history, with generic diversification among extant lineages pre-dating the Oligocene (> 33 Ma). Ancestral-area reconstructions indicate the presence of brevicipitids in the EABR since the Oligocene, and support a scenario of palaeoendemics surviving in EABR refugia. Ancestral-area reconstructions indicate that the central Eastern Arc Mountains (EAM) formed the initial centre of diversification of forest brevicipitids. Measures of PD show that diversity varies across the EABR but is highest in the EAM. Constant net diversification rate in brevicipitids is a significantly better fit than alternative, rate-variable models.Main conclusionsThe degree of persistence of forest habitats appears to be a contributing factor to the varying levels of diversity across the EABR in brevicipitids (and other organisms). In contrast to the Southern Highlands and Ethiopian Bale Mountains, the EAM stands out as an area that enabled the constant accumulation of brevicipitid species over a long period of time.
A new species of dwarf chameleon is described from a restricted patch of relict indigenous forest in KwaZulu-Natal (KZN), South Africa. Its specific status is confirmed by phylogenetic analyses using molecular markers (1390 bp of mitochondrial 16S and ND2). The node defining this species is highly supported with both analyses (100% posterior probability, 100% bootstrap support). This species (Bradypodion ngomeense sp. nov.) is part of a larger clade of forest species, but is itself confined to a single forest (Ngome Forest). The molecular patterns of other forest restricted species in KZN were examined with a view to elucidate their patterns of distribution. It is postulated that these patterns may be the result of climatic shifts during the Pleistocene on the extent of forest cover which afforded multiple contact opportunities between coastal and montane forest elements with possible opportunities for gene flow between forests. At present, these patches are under threat from human activities such as forest resource extraction, sugar cane and pine plantations. Their small distributions and possibility for future habitat loss make these species of conservation concern. The taxonomic status of a recently described species Bradypodion nkandlae (Raw & Brothers 2008) is found to be conspecific with B. nemorale and is herewith synonymised.
The taxonomic history and composition of the genus Bradypodion as construed by Klaver & Böhme (1986) and new morphological and molecular data relevant to the taxonomy of the group is reviewed. The combined evidence strongly supports a formal rearrangement of the group into three distinct genera. Bradypodion, type species Chamaeleo pumilus Daudin 1802, is retained for the southern African species. Two new genera are erected to accommodate additional well-diagnosed clades within central and east African species previously referred to Bradypodion. Species of the “fischeri complex” are assigned to Kinyongia gen. nova, whilst the endemic Mulanje chameleon is placed in the monotypic genus Nadzikambia gen. nova.
We examined approximately 600 specimens that represent the Praomys delectorum species group (Muridae: Murinae: Praomyini), a rodent complex restricted to Afromontane landscapes in East Africa and currently viewed as a single species. Morphometric analyses of 21 population samples consistently disclosed cohesive patterns of craniodental differentiation that support the recognition of three species: Praomys delectorum Thomas, confined to extreme southern Malawi; P. melanotus Allen & Loveridge, found in highlands of south-western Tanzania and contiguous northern Malawi; and P. taitae Heller (including octomastis Hatt), distributed in mountains and foothills of southern Kenya and northern and central Tanzania. Populations of the P. delectorum group are patchily distributed in moist montane forest, most collecting localities falling within 1000–2400 m, and their range collectively coincides with the Tanganyika–Nyasa Montane Forest Group sensu Moreau. Patterns of faunal similarity derived from distributions of 65 species of terrestrial small mammals recorded from Tanzania's highlands, including the Eastern Arc Mountains, demonstrated pronounced geographical discontinuities in montane associations but failed to uncover a prominent vicariant role for the Makambako Gap. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165, 420–469.