Zoosyst. Evol. 97 (1) 2021, 161-179 | DO! 10.3897/zse.97.57297 gee BERLIN A new chameleon of the 7rioceros affinis species complex (Squamata, Chamaeleonidae) from Ethiopia Thore Koppetsch!, Petr Neéas*, Benjamin Wipfler? 1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113, Bonn, Germany 2 Archaius, Rozdélovaci 1380, CZ-66434, Kurim, Czech Republic http://zoobank.org/COEFA214-ACB8-45BD-B763-822C41FEEDB6 Corresponding author: Thore Koppetsch (t.koppetsch@leibniz-zfmk.de) Academic editor: Johannes Penner # Received 5 August 2020 Accepted 18 February 2021 @ Published 10 March 2021 Abstract A new species of chameleon, 7rioceros 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 morpho- logical characteristics, phylogenetic discordances of previous studies and biogeographical patterns, this new species is assigned to the Trioceros affinis (Ruppell, 1845) species complex. An updated comprehensive key to the Trioceros found in Ethiopia is provided. Key Words Bale Mountains, biogeography, Ethiopia, Great Rift Valley, key, morphology, new species, systematics, taxonomy, Trioceros wolfgangboehmei sp. nov. Introduction Chameleons can be considered as one of the most fasci- nating groups of squamate reptiles, not only due to their exceptional locomotion, behaviour or anatomical adap- tations, but also because of their extraordinary species diversity and radiations (Raxworthy et al. 2002; Tolley et al. 2008, 2011). The more than 215 described species of the family Chamaeleonidae are distributed from Africa (including Madagascar), Southern Europe and the Middle East to parts of South Asia (Neéas 2004; Tilbury 2010; Uetz et al. 2020). Apart from the fact that Madagascar is a well-known hotspot of chameleon diversity (Townsend et al. 2009; Gehring et al. 2012; Prétzel et al. 2018), it was hypothesised that this family originated in mainland Africa (Tolley et al. 2013; Cerfansky et al. 2020). The African continent harbours not only a broad variety of morphologically and ecologically distinct chameleon species, but, in particular, also a high number of montane endemics that are restricted to single moun- tains or mountain ranges. Examples include the genus Kinyongia (Menegon et al. 2002, 2009; Ne€as et al. 2009; Hughes et al. 2017), but also several recently described species of the genus Trioceros like T. hanangensis (Mt. Hanang) from Tanzania (Krause and Bohme 2010) or 7! kinangopensis (Kinangop Peak, Aberdare Mountains), T. narraioca (Mt. Kulal), 7. ntunte (Mt. Nyiru) and T. ny- irit (Mtelo Massif) from Kenya (Neéas et al. 2003, 2005; Stipala et al. 2011, 2012). This Eastern Afromontane region (EAR) exhibits an outstanding level of species richness, endemism and di- versification (Mittermeier et al. 2011; Demos et al. 2014; Mairal et al. 2017) and can be also considered an ex- traordinary biogeographical unit for chameleon diversity (Mariaux and Tilbury 2006; Ceccarelli et al. 2014; Men- egon et al. 2015; Hughes et al. 2018). The geographical Copyright Thore Koppetsch et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 162 uniqueness of this region 1s shaped by the Great Rift Val- ley, a geological longitudinal split between the Somalian and Nubian Plate, running nearly 5,000 km through the African Plate from the Red Sea to Mozambique (Chorow- icz 2005). These topographical and, consequently, bioge- ographical patterns are not only present along the tropi- cal East African Great Rift in Tanzania or Kenya, where many endemic TJrioceros species live. They can also be found in the Ethiopian Highlands, another biodiversity hotspot at the Horn of Africa (Mittermeier et al. 2011). This northern part of the Great Rift System is located south of the Afar Depression between the Abyssinian and the Harrar Massif within the Ethiopian Highland Plateau. Species differentiation is also common in this part of the Rift and numerous examples of plants, insects and verte- brates have been described tn the past (Mairal et al. 2017; Manthey et al. 2017). Particularly, for several mammals and amphibians, e.g. for the Ethiopian wolf (Gottelli et al. 2004), the gelada baboon (Belay and Mori 2006) or anurans (Evans et al. 2011; Freilich et al. 2014, 2016), phylogeo- graphic disyunctions across the Ethiopian Rift were shown. The Ethiopian Chameleon Trioceros affinis (Ruppell, 1845) is a small chameleon species, endemic to the Ethi- opian highlands, where it is widely distributed in altitudes higher than 1,600 m above sea level (Neéas 2004; Largen and Spawls 2010). However, its distribution range within the Ethiopian region includes multiple populations isolat- ed in high-altitude forest patches (Ceccarelli et al. 2014). Since some of these populations are particularly isolated and additionally located both west and east of the Ethio- pian Rift, the separation by this geological trench might have acted as a long-term dispersal barrier as it also does in other taxa (Mairal et al. 2017). This variation of 7° af- finis populations across the Ethiopian Highlands was al- ready indicated by Neumann (1905) based on differences in the shape of the cranial crests and granularity of the body scalation. Neéas (1994, 2004) stressed the possible existence of various distinct and geographically separated forms currently assigned to T affinis as well. Subsequent- ly, it was shown that considerable genetic divergence be- tween western and eastern populations separated by the Rift is present and that 7° affinis actually should be re- ferred to as a species complex (Ceccarelli et al. 2014). Despite limited sampling, Ceccarelli et al. (2014) were able to identify two lineages, one west of the Rift around Addis Abeba and another one based on material from Goba and Dinsho, which diverged during the Pliocene (4 mya) and suggested their distinctness at species level. Ruppell (1845) separated Trioceros affinis from Cha- maeleo senegalensis based on the lack of a gular crest and referred to ‘Abyssinia’ as a type locality. This refer- ence to an uncertain and broad locality description may not reflect the actual diversity and divergence of this chameleon. Unfortunately, it is not possible to narrow down the type locality based on further or more detailed locality data from other herpetological examinations of Ruppell’s material and descriptions (von Heyden 1827). In later treatises by Gray (1864) only additional morpho- logical characteristics of T° affinis are described but no zse.pensoft.net Koppetsch, T. et al.: A new chameleon from Ethiopia specific remarks concerning its distribution are given. Fi- nally, when assigning lectotype material for this species (Fig. 1), Mertens (1967) also had no evidence for further concrete specifications of the locality likewise and had to refer to the vague and imprecise term ‘Abyssinia’ again. Considering the shown genetic discordance and possible additional taxonomic assessments, a restriction of the type locality for 7 affinis might be possible in future. In the context of northern Eastern Afromontane bio- diversity hotspots, the Bale Mountains are an extraordi- nary biogeographical unit within the Ethiopian Highlands (Hillmann 1988). This extensive south-central Ethiopian high-elevation plateau and massif above 3,000 m is a re- markable centre of endemism: Numerous endemic species are restricted to this area, e.g. several rodents, like Arvi- canthis blicki, Dendromus lovati, Lophuromys melanon- yx or Stenocephalemys albocaudata (Lavrenchenko et al. 1997), frogs, like Altiphrynoides malcolmi, Balebreviceps hillmani, Ericabatrachus baleensis or Spinophrynoides osgoodi (Largen and Spawls 2011; Gower et al. 2013), or the Bale Mountains Adder Bitis harenna (Gower et al. 2016), and also the endangered Mountain Nyala 7ragela- phus buxtoni occurs in this area (Refera and Bekele 2004). Two Trioceros species endemic to this area are already known: T. balebicornutus (Tilbury 1998) and 7: harennae (Largen 1995). In specimens assigned to 7: affinis from the museum collections of the Forschungsinstitut und Natur- museum Senckenberg, Frankfurt, Germany (SMF) and the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK), individuals from northern re- gions of the Bale Mountains could be identified by the first author. Examination of collection material from this centre of endemism on the one hand, and in biogeograph- ical separation across the Ethiopian Rift from other pop- ulations, on the other, revealed the presence of a cryptic species within the 7rioceros affinis species complex in this region. In the following, we describe it as a new spe- cies and provide a re-description of the lectotype material. Material and methods Morphological analysis We examined a total of 66 specimens preserved in 70% ethanol obtained from the museum collections of the For- schungsinstitut und Naturmuseum Senckenberg, Frankfurt, Germany (SMF) and the Zoologisches Forschungsmuse- um Alexander Koenig, Bonn, Germany (ZFMK). Scale counts were made using a stereo microscope and morphological measurements were taken with a vernier caliper to the nearest 0.1 mm. Morphometric features for the morphological analyses were selected based on previ- ous taxonomic studies of chameleons (Neéas 1994; Necas et al. 2005; Barej et al. 2010; Stipala et al. 2011; Green- baum et al. 2012). In addition to these meristic and quan- titative mensural characters, we added three qualitative morphological features in order to account for the pres- ence and characterisation of the prominent, distinctive Zoosyst. Evol. 97 (1) 2021, 161-179 patterns of the head, body and dorsal crest scalation. These are the enlarged dorsal crest scales (EDCS), het- erogeneous body scalation (HBS) and rugose head scala- tion (RHS) (detailed explanations provided below). For all specimens examined, we determined sex and meas- ured the following 26 morphometric features (see Table 1 and Suppl. material 2: Individual mensural and meristic measurements): SVL — snout-vent length, from the snout tip to the frontal margin of the cloacal fissure; TaL — tail- length, from the frontal margin of the cloacal fissure to the tail tip; TL — total length as a sum of SVL + TaL; HL — head length from superior tip of casque to snout tip; H'W — head width, measured at widest point just posterior to eyes; ML — mouth length, from tip of rostral to jaw com- missure; CH — casque height, from commissure of jaw to superior tip of casque; CN — casque exceeding neck, dis- tance between top of casque and level of the dorsal crest on neck; CE — casque-eye length, measured diagonally from posterior margin of orbit to superior tip of casque; SL — snout length, from tip of snout to anterior margin of orbit, ED — eye diameter, measured horizontally at cen- tre of eye; IOS — number of inter-orbital scales, number of scales between supra-orbital crests, including crests scales; PCL — length of parietal crest, from superior tip of casque to rostral border of parietal crest; CC — cranial crest gap, measured across the crown between raised su- praorbital crests at mid-eye; IL — inter-limb length, from axillary to inguina; FLL — forelimb length, from elbow to wrist; LHLL — lower hindlimb length, from knee to heel; UL — number of supralabials to posterior margin of orbit; LL — infralabials to posterior margin of orbit; SEH — number of scales between eye and posterior margin of head; FSM -— flank scales at midbody; SDC — enlarged (more than twice the size of the surrounding dorsolater- al scales) and conical scales forming dorsal crest, from head to frontal margin of the cloacal fissure; SCP — scales forming parietal crest; EDCS — presence of significant- ly enlarged (more than twice the size of the surrounding dorsolateral scales) conical dorsal crest scales; HBS — presence of a prominent heterogeneous body scalation; RHS — presence of a rugose head scalation consisting of enlarged (more than 1.2 times the size of the surrounding dorsolateral scales) and slightly conical scales. The following eight ratios were calculated: TaL/SVL — tail length/snout-vent length ratio; RTL — relative tail length, tail length/total length ratio; HW/HL — head width/head length ratio; RCH — relative casque height, CH/ML; RCN — relative length of casque exceeding neck, CN/ML; ED/HW -— eye diameter/head width ratio; FLL/ SVL — forearm length/snout-vent length ratio; LHLL/ SVL — lower hindlimb length/snout-vent length ratio. Furthermore, colouration and patterns were recorded for the preserved individuals examined (see Suppl. mate- rial 2: Individual mensural and meristic measurements). For obtaining information on hemipenial morphol- ogy, hemipenes of Trioceros affinis (ZFMK 54264) (Fig. 4A—C) and Trioceros sp. nov. (ZFMK 84811, holo- type) (Fig. 4D—-F) were scanned with a Skyscan 1173 u-computer-tomograph (u-CT) (Bruker, Billerica / USA) 163 at the Museum Koenig in Bonn Germany. Hemipenes preparation was modified after Ziegler and Bohme (1997) and Zaher (1999): For each specimen the already everted left hemipenis was removed, submerged in a solution of 3% KOH for around 15 minutes until it became flexible and finally inflated with coloured petroleum jelly. For increasing X-ray absorbance, all samples were stained with iodine for two days following Protzel et al. (2015). The scans were performed with the following parame- ters: 30 kV for Trioceros affinis and 31 kV for Trioceros sp. nov., 133 uA, 1000 ms exposure time for Trioceros affinis and 1100 for Trioceros sp. nov., 0.3° rotation steps over 360° for Trioceros affinis and 0.35° over 360° for Trioceros sp. nov., frame averaging of 7 for Trioceros affinis and 8 for Trioceros sp. nov., random movement of 15, image pixel size 8.1627 um for Trioceros affinis and 6.0343 for 7rioceros sp. nov. Both scans are deposit- ed in the collection of the Museum Koenig in Bonn and can be downloaded at Morphobank (https://morphobank. org) under project number P3948. The scans were recon- structed with the software NRecon (Bruker, Billerica / USA). Segmentation of the resulting scans was per- formed with Amira 5.3 (Thermofisher, Waltham / USA). Subsequent volume rendering was done with VG Studio 3.3.4 (Volume Graphics, Heidelberg / Germany). Final images and plates were edited with Adobe CS6 (Ado- be, San Jose / USA). Terminology describing hemipenial structures follows Klaver and Bohme (1986) and Protzel et al. (2015, 2018). Principal component analysis of morphological data A principal component analysis (PCA) of chosen men- sural and meristic characters (indicated in Table 1; ratios and TL as a cumulative character were not included in the principal component analysis in order to avoid the dou- ble use of characters and resulting bias) was conducted using the Ade4 package (Dray and Dufour 2007) for the software R (R Development Core Team 2012). Regres- sion residuals of continuous variables were calculated. The analysis was performed by using mixed variables. We applied the Broken Stick analysis using the “evplot” function from the package “Imom” in R to calculate the number of the meaningful components. Based on both quantitative and categorical variables, the first two prin- cipal components (PCs) are visualised in Fig. 5. We applied a non-parametric MANOVA (Per-MANO- VA) on the first 3 principal components (obtained through Broken Stick analysis) using the software package PAST (Hammer et al. 2001) tn order to investigate if the specimens from the northern Bale region (Goba and Dinsho) signifi- cantly differ from those of other examined populations of Trioceros affinis . We tested our meaningful components for normal distribution, however since there are more depend- ent variables than sampling units included in the analysis, we used the non-parametric MANOVA test (Per-Manova) over the parametric one (Anderson 2001; Kean et al. 2015). zse.pensoft.net 164 Results Re-description of the lectotype of Trioceros affinis (Riippell, 1845) Systematics Chamaeleonidae Werner, 1902 Trioceros Swainson, 1839 Trioceros affinis (Riippell, 1845) Material examined. Lectotype: SMF 16402, adult male; origin: ‘Abyssinia’; collected by E. Ruippell, 1834; desig- nated by Mertens (1967) (Fig. 1A—C). Paralectotype: SMF 16403, adult female, same collec- tion data as the lectotype (Fig. 1D—F). Re-description of the lectotype. The adult male lecto- type (SMF 16402) of T7rioceros affinis is a small-sized and slender chameleon without outstanding ornamen- tation on the body and head. It has a total length of 148.3 mm (snout-vent length of 68.9 mm and a tail length of 79.4 mm) (Fig. 1A). The tail is slightly longer than the rest of the body (RTL 0.54), heavily thickened at the base due to the presence of hemipenes situated in their pockets. The extremities are thin, the hind legs are slight- ly wider in diameter but equal in length. The head is relatively short and blunt (HW/HL 0.52) with a low casque (RCH 0.85) merely exceeding the lev- el of the neck (RCN 0.2). The head surface is covered with slightly enlarged flattened scales, which are more convex and slightly rugose between the canthi rostrales (Fig. 1B, C; see Suppl. material 3: Dorsal head views). The head crests, typical for the genus 7rioceros, are well developed and ornamented with slightly enlarged, convex scales. The canthus lateralis is well developed in all its parts. It starts with two paramesial scales on each side of the head, builds a margin of the low casque, continues as slightly less expressed supra-ocular crest with a more warty appearance on the canthus rostralis, which conjoin just above the tip of the snout forming a low ridge. The parietal crest consists of 5 scales merely larger than the surrounding ones, building a faint yet sharp medial ridge on the top of the casque. The temporal crest, consisting of 6 enlarged scales 1s expressed as a lateroventral emargina- tion of a rugose triangular field below the lateral crest just behind the orbit and posterodorsally from it. It reaches the canthus lateralis proprius at about half of 1ts length where it fuses with it. There are 16 upper labials and 16 lower la- bials on both sides of the head. The eye turrets are covered with a granular homogeneous scalation. The nostril is sit- uated at half distance between the tip of the snout and ros- troventral margin of the orbit. The body is covered with subhomogenous scalation, the standard scales are convex, not flat, of sub-circular or sub-oval form and of approx- imately same size all over the body, tail and extremities, with a tendency to become smaller ventrally towards the midventral line and on the distal part of the tail towards zse.pensoft.net Koppetsch, T. et al.: A new chameleon from Ethiopia the tail tip. On the flanks, they are ordered in more or less longitudinal or diagonal long fields (running in craniodor- sal-caudoventral orientation) that are grouped in rows of two (rarely three) scales. Here no interstitial skin is ex- posed, with the exception of the gular region, where later- oventrally, an inconspicuous groove can be observed be- tween the slightly enlarged, spindle-shaped scales. There is one irregular, interrupted line of 8 significantly enlarged lenticular scales (the largest ones are about 3 times longer and 2 times wider than the surrounding standard scales). These scales are separated from each other by 3—7 stand- ard scales, oriented longitudinally at about 2/3 of the body height and range from the shoulder to the pelvic region. Anteriorly, the dorsal crest runs posteriorly of the casque, extending as a continuous well-developed, but low, crest consisting of conical scales of about double width and double height compared to the surrounding standard scales. The dorsal crest reaches in its described form the level of the groin, decreasing in height and formed by smaller subconical scales up to the first 1/4 of the tail with a more inconspicuous appearance. Dorsolat- erally on both flanks, the first line of scales bordering the dorsal crest is slightly enlarged. The gular crest is absent, instead, a rather narrow tri- angular field scattered with significantly smaller granular scales is present and ranges from the mentum to the arch of the hyoid. The ventral crest consists of a slightly en- larged, midventral line of scales ranging from the shoul- der to the frontal margin of the cloacal fissure, which is separated into two parallel rows just along the umbilical scar, and ends at the ventral part of the tail. Detailed meristic and mensural measurements of the lectotype are given in Table 1 and in the Suppl. material 2: Individual mensural and meristic measurements. Coloration in preservative. The animal is more or less uniformly greyish beige all over the body without any con- Spicuous pattern. Only a slight yellow hue can be found midventrally on the body, tail and the soles. The top of the head is slightly brownish. The claws are brownish yellow. Some prominent scales in the head region have a blackish centre, probably as a result of a mechanical bruise. Variation. The adult gravid female paralectotype (SMF 16403) (Fig. 1D-—-F) shows only minor morphologi- cal differences compared to the male lectotype. It has a lower total length (TL 146.0 mm) and longer tail (RTL 0.50). Also, it has a relatively higher casque (RCH 0.89) (Fig. 1E, F). Both for the upper and lower labials 19 are present. 56 flank scales at midbody are present (vs. 54 in the lectotype) and a slightly lower number of scales forming the dorsal crest can be found (49 vs. 53 in the lectotype). The canthus parietalis 1s formed by only 3 scales. In contrast to the lectotype, the entire head region is coloured whitish without any brownish hue. Detailed meristic and mensural measurements of the pa- ralectotype are given in Table 1 and in the Suppl. material 2. Based on our comparative examination of 7) affinis specimens from different regions of Ethiopia, the lecto- 165 Zoosyst. Evol. 97 (1) 2021, 161-179 SS, ‘~ x Org, Ose: My : ‘ se i i Figure 1. The preserved male lectotype (SMF 16402) (A.) and female paralectotype (SMF 16403) (B.) of Trioceros affinis collected Head of the lectotype in left (C.) and right (D.) view and Morris Flecks. F.) view. Scale bars represent 1 cm. Photos by Robert Mertens. Head morphology: by E. Ruppell 1834 and designated by of the paralectotype in left (E.) and right ( lectotype material. Compared to the lectotype material, type material can be clearly assigned to populations west of the Ethiopian Rift. Individuals from south-western single individuals from Addis Abeba showed a more het- erogeneous scalation on parts of the flanks by having sin- Ethiopia showed a lower body length compared with the zse.pensoft.net 166 gle lateral scales that are enlarged more than 1.5 times compared to the surrounding scales. However, this het- erogeneous scalation is not as prominent and extensive as in individuals of the northern Bale region. Actually, the latter were clearly distinct from other 7! affinis spec- imens, which in the following is shown by significant statistical support and conspicuous differences in other diagnostic characters. Justification for a new species of Trioceros based on statistical analyses. According to our PCA analysis based on the morphological examination of the lectotype mate- rial of T. affinis as well as T: cf. affinis specimens from various localities in Ethiopia the first three components (those obtained through Broken Stick analysis) explained 24%, 17%, 10% of the total variation respectively). The first component was clearly discriminative between the species (Fig. 5; see Suppl. material 4: PCA Axis 1 vs. Axis 3 and PCA Axis 2 vs. Axis 3). The highest load- ings for the first component referred to the continuous variables SDC, PEL, SCP and the categorical variables EDCS, HBS and RHS (Suppl. material 1: PCA loadings). Based on this component the new species has a lower number of SDC, a longer PEL and a higher number of SCP than 7: affinis. Also, the new species has the exclu- sive characters EDCS, HBS and RHS, while none of the 7. affinis individuals did. The first axis did not separate different sexes in any of the species. The lectotype and paralectotype of 7: affinis show no significant differences to individuals of other populations included in our statis- tical analyses. Even though only the first principal com- ponent showed clear shape separation among the studied species, we still used all 3 meaningful components in the Per-MANOVA analysis. Based on Per-MANOVA the two species were significantly different from each other (F = 43.7, p = 0.0001). According to the extensive morphological investigation of both the lectotype material of Trioceros affinis sensu stricto and individuals of 7: affinis from other locations in Ethiopia, and the significant statistical distinctness of our new species from all the other 7: affinis, we here formally describe T7rioceros wolfgangboehmei sp. nov. Systematics Chamaeleonidae Werner, 1902 Trioceros Swainson, 1839 Trioceros wolfgangboehmei, sp. nov. http://zoobank.org/20A 1 D4E2-40E5-49C 1-8F4D-1580EAD87531 Suggested common English name: Wolfgang Bohme’s Ethiopian Cha- meleon Material examined. Holotype: ZFMK 84811, adult male, Dinsho (3,130 m as.l. / 7°06'10"N, 39°47'25"E), Bale Mountains, Ethiopia, collected by Petr Neéas on the 6. August, 2004 (Fig. 2A, C, D). zse.pensoft.net Koppetsch, T. et al.: A new chameleon from Ethiopia Paratypes: ZFMK 84812, adult female, with the same locality and collecting data as the holotype; ZFMK 84813, adult female, Goba (2,740 m a.s.l. / 7°00'36"N, 39°57'28"E), Bale Mountains, Ethiopia collected by Petr Neéas on the 7. August, 2004 (Fig. 2B, E, F); ZFMK 63063, adult female, 10 km from Goba (2700 m a.s.1.), Bale Mountains, Ethiopia collected by Colin Tilbury in October 1996. Diagnosis. Trioceros wolfgangboehmei sp. nov. is a small-sized chameleon of the T7rioceros affinis species complex (sensu Ceccarelli et al. 2014). It can be distin- guished from all other members of the same species com- plex by the following combination of characters: (1) presence of a prominent and well-developed dor- sal crest consisting of a relatively low number of significantly pointed and enlarged conical scales, forming a single row and reaching along the ante- rior half the tail; (2) top of the casque posteriorly raised above the dorsal crest; (3) heterogeneous body scalation with both small scat- tered standard scales and enlarged flattened plate- like scales; (4) long canthus parietalis formed by 9-12 slightly en- larged scales; (5) rugose head scalation consisting of enlarged scales forming the cranial crests that fill the area between the lateral and temporal crest and the posterior rim of the orbit; (6) relatively high number of flank scales at midbody (53-59); (7) relatively short snout-vent length (up to 66 mm); (8) aunique hemipenial morphology including shallow calyces with smooth margins on the truncus, four pairs of thick, pointed and thorn-like papillae and two pairs of non-serrated rotulae. Description of the holotype. The adult male holotype (ZFMK 84811) of Trioceros wolfgangboehmei sp. nov. is a small-sized chameleon with a total length of 156.3 mm (snout-vent length of 65.3 mm and a tail length of 91.0 mm) (Fig. 2A). The head is relatively short, 18.9 mm long (HW/HL 0.54). The head scalation is rugose con- sisting of enlarged scales forming the cranial crests and filling the area between the lateral and temporal crest and the posterior rim of the orbit (Fig. 2C, D; see Suppl. material 3: Dorsal head views). The top of the casque is raised posteriorly above the dorsal crest. Nine convex, enlarged and tubercular scales form the parietal crest. The casque is 11.7 mm high (RCH 0.91) and exceeds the dor- sal crest on the neck by 3.2 mm (RCN 0.25). The parietal crest is 9.2 mm long. The temporal region is covered by prominent enlarged scales of larger size than the standard scales on flanks and limbs. The gular crest is absent in preserved specimens — in life, it is visible as two parallel paramesial skin folds on the throat. The temporal crest is Zoosyst. Evol. 97 (1) 2021, 161-179 167 Table 1. Mensural and meristic measurements of type specimens of 7rioceros wolfgangboehmei sp. nov. and male, female and juvenile individuals of 7: affinis examined (including the lectotype material). For the specimens of 7: affinis linear measurements (in mm) and scale counts are given as mean values (Mean) + standard deviation (SD) and sample size (N), minimum (Min) and maximum (Max) are shown. Presence (present) and absence (—) of characters are indicated. Characters used for statistical analysis are marked with a star (*). See Materials and Methods for explanation of the single character abbreviations. Species Trioceros wolfgangboehmei sp. nov. Trioceros affinis Sex male female female female male female male female juvenile Locality Dinsho Dinsho Goba Goba_ Abyssinia Abyssinia Ethiopia Ethiopia Ethiopia holotype paratype paratype paratype lectotype paralectotype N = 26 N = 32 N=4 ZFMK ZFMK ZFMK = ZFMK SMF SMF Meant Min Max Meant Min Max Mean+ Min Max 84811 84812 84813 63063 16402 16403 SD SD SD SVL* 65.3 65.8 60.3 oh evs 68.9 f3.3 74.73 + 67.6 846 77.954 63 88.3 51.88+ 48.2 55.2 4.57 6.31 2.87 TaL* 91.0 81.4 72.0 eS 79.4 72.7 89.87+ 77.2 110.2 80.334 62.1 92.4 60.83+ 58.1 62.5 7.66 7.41 1.90 aL 156.3 147.2 132:3 137 148.3 146.0 164.60 + 145.8 183.1 158.28+ 133.2 178.4 112.70 106.3 116.4 10.50 12.46 +441 ALS 18.9 20.3 17.9 18.1 18.8 20.2 20.37 + 18. 22:7 20:70. # 18:2) 22-7 V5:90:+) 15:4 16:8 125 1.25 0.62 HW* 10.2 10.8 9.1 OLD 9.7 10.6 TO05:+° -815° 13:9 T10@O:F. 8:2. <11:9- 66:43 8 G21 113 0.88 0.48 ML* 12.9 15.4 12.7 12.4 13.6 14.8 1462+ 135 164 15.194 TS) AZ ABO =i 11.8 0.81 1.26 0.38 CH* 11.7 12.6 12.2 11.2 11.6 algoyal 12.27+ 10.2 142 12.764 11 141 963+ 89 10.5 0.96 0.91 0.67 CN* 302 4.6 4.1 3.0 2.7 333 3.17 + 2.2 48 3.21 + O53) @ASO? A220 AG OF 0.56 0.59 0.35 CE* 8.3 8.8 8.4 8.9 8.3 9.4 8.88 + 6.5 108 877+ 6.9 104 648+ 5.7 6.9 1.00 0.81 0.53 Sie 5.4 6.3 5.0 a7. 6.3 7.6 6.48 + oh Se af 6.98 + 5.2 15.7 460+ 3.9 5 0.61 1.89 0.48 ED* 6.1 5.8 5.2 4.9 6.5 5.4 6.50 + 5 7A 6.56 + Bye 725 6 660k 4:9 "6.1 0.53 0.45 0.56 lOS* 12 11 11 11 13 12 13.08 + 11 15 13.23 + 11 15: “h2:50%, 12 13 1.02 Le] 0.58 PCL* 9.2 12.6 10.1 Or2 4.1 38 4.81+ 3.1 79 4.60 + 24 76 3.604 3 4.8 1.14 0.94 0.83 cCc* 8.5 8.4 7.6 7.6 7.6 8.1 8.34 + 74 94 8533:-+ fl 9.9 665+ 6 Es 0.59 0.74 0:53 [es 39.2 36.4 34.1 31.9 37.8 43.9 41.10+ 346 49 45.07+ 34.9 55.2 27.354 224 30 3:70 4.99 3.40 FLL* 11.9 11.8 10.7 9.5 14.9 15.4 13.64+ 11.1 16.2 1360+ 11.2 16.8 1053+ 98 11.5 1.37 1.46 0.71 LHLL* 10.8 11.4 ay Ter 14.0 15.8 1336): 10:4 -15 13736 « «hls? 16.9 -10i05:= --933 ~AL:2 1.10 1.25 0.83 UL* 16 16 15 16 14 15 16.27 + lS 19 16.62 + 13 19 1450+ 13 16 1.64 1.60 1.29 LL* 17 15 15 15 14 15 15.27 + 12 17 14.92 + yes 18 15.00+ 14 16 1.12 1.26 1.15 SEH* 6 7 ip i: 7 7 8.00 + 6 11 Salo st 6 10 6.50+ 6 7 1.17 0.97 0.58 FSM* 53 58 59 53 54 56 60.154 48 69 62.69 + 56 69.~ -b2-50: » 51 54 5.30 4.03 1.29 SDC* 37 33 35 38 53 49 62.314 47 78 64.38 + 56 76 49.75+ 46 58 8.00 5.07 5.68 SCP* 10 12 9 11 5 3 442+ 3 7 4.35+ 1 6 ony soln 3 5 1.14 1.16 0.96 EDCS* present present present present - - - - - - - - - - - HBS* present present present present - - - - - - - - - - - RHS* present present present present - - - - - - - - - - - TaL/ 1.39 1.24 aks) 1.29 1.15 0.99 Lee 2OTE e 1:O7© &T: 5: LOSE. 10:87 wel ed fe Lab. @ la) SVL 0.10 0.08 0.05 RTL 0.58 0.55 0.54 0.56 0.54 0.50 0.554 0.52 0.60 0514+ 047 054 054+ 053 0.55 0.02 0.02 0.01 HW/HL (0.54 0.53 0.51 0.52 0.52 0.52 049+ 0.45 066 049+ 044 053 053+ 0.51 0.54 0.05 0.02 0.01 RCH 0.91 0.82 0.96 0.90 0.85 0.89 0.844 0.71 0.99 084+ 0.70 0.96 085+ 0.75 0.92 0.07 0.06 0.07 RCN 0.25 0.30 0.32 0.24 0.20 0.22 0.22+ 0.15 0.34 O.214 0.16 030 O0.20+ 0.16 0.25 0.04 0.04 0.04 ED/HW 0.60 0.54 0.57 0.52 0.67 0.51 O:65-) 0:53 0:79. Or6b:+ —0:55 O76 O0:66s% O60 OL71 0.07 0.06 0.05 FLL/ 0.18 0.18 0.18 0.16 0.22 0.21 O.lS:+ -Ocl5) O:22~ Oreck