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Acanthocercus ceriacoi MARQUES, PARRINHA, SANTOS, BANDEIRA, BUTLER, SOUSA, BAUER & WAGNER, 2022

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Ceríaco’s Tree Agama
P: Sardão Arborícola de Ceríaco 
SynonymAcanthocercus ceriacoi MARQUES, PARRINHA, SANTOS, BANDEIRA, BUTLER, SOUSA, BAUER & WAGNER 2022
Stellio nigricollis BOCAGE 1866: 43 [lapsus calami; see WAGNER et al. 2018]
Stellio atricollis — PETERS 1881: 147 [part]
Stellio atricollis — BOCAGE 1895: 22) [part]
Agama atricollis — BOULENGER 1885: 256 [part]
Agama atricollis — BOULENGER 1905: 110 [part]
Agama atricollis — FERREIRA 1903: 15 [part]
Agama atricollis — MONARD 1937: 58 [part] 
DistributionC/NW Angola (from central Malanje to Bengo Province in the west and Uíge Province in the north).

Type locality: Cangandala National Park headquarters [-9.819417° N, 16.653861° E, 1089 m a.s.l.], Malanje Province, Angola.  
Reproduction 
TypesHolotype: CAS 258430, adult male collected by Mariana P. Marques, Luis M.P. Ceríaco, Suzana A. Bandeira, Edward L. Stanley and Jens V. Vindum on 14 September 2015.
Paratypes: CAS 2584295; CAS 258431; CAS 258433; CAS 258434; MHNCUP/ REP 860 
DiagnosisDiagnosis: Segmentation of the tail into distinct whorls places the new species in the genus Acanthocercus. Within this genus, heterogeneous body scalation refers it to the cyanogaster/atricollis group and the high density of enlarged trunk scales, in combination with a black patch on the shoulder, place it in the A. atricollis complex. Acanthocercus ceriacoi is a medium sized lizard (mean SVL 122.5 mm [99–143 mm]; mean total length 285 mm [224–330 mm]) for its genus and overall is very similar in morphology to A. cyanocephalus. However, it may be distinguished from all other species of the complex by its meristic characters. The new species has lower scale counts when compared to other species of the A. atricollis complex group (mean scale rows around midbody 108, mean dorsal scale rows 64, mean ventral scale rows 85). In life, displaying males are characterized by a blue coloration of the head; neck, shoulders, body and limbs brownish with a pattern of white and black stripes and speckles; and a pale vertebral band from the neck to the tail, distinct anteriorly between the flanks, becoming more indistinct along a brown background posteriorly. (Marques et al. 2022)

Comparisons: Acanthocercus ceriacoi can be distinguished from other taxa of the A. atricollis complex by the following characters:
(a) Acanthocercus atricollis has a larger SVL (mean 129 mm vs. 123 mm) and total length (mean 302 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (112 vs. 108), dorsal scales (67 vs. 63), and ventral scales (87 vs. 85) than A. ceriacoi. The mean number of precloacal pores is higher in A. atricollis than in A. ceriacoi (22 vs 18). Acanthocercus atricollis presents a blue or green head and ventral surface, in some individuals extending to the body, while in A. ceriacoi sp. nov. the blue is limited to the head.
(b) Acanthocercus branchi has a smaller SVL (mean 109 mm vs. 123 mm) and total length (mean 262 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (117 vs. 108), dorsal scales (73 vs. 63), and ventral scales (88 vs. 85) than A. ceriacoi. The ranges of these characters largely overlap in both species, although they are higher in A. branchi than in A. ceriacoi. The mean number of precloacal pores is higher in A. branchi than in A. ceriacoi (20 vs 18).
(c) Acanthocercus cyanocephalus has a larger SVL (mean 138 mm vs. 123 mm) and total length (mean 334 mm vs. 285 mm). It is morphologically very similar to A. ceriacoi and its meristic differences are subtle. Acanthocercus cyanocephalus has a higher mean dorsal scale count (68 vs. 63), and a lower mean number of ventral scales (80 vs. 85) than A. ceriacoi sp. nov.. The mean number of precloacal pores is higher in A. cyanocephalus than in A. ceriacoi (22 vs 18). The major difference between these two species is the coloration of displaying males, which in A. cyanocephalus is characterized by the blue coloration of the head extending to the neck and shoulders, whereas in A. ceriacoi the blue is limited to the head.
(d) Acanthocercus cyanogaster has a smaller SVL (mean 101 mm vs. 123 mm) and total length (mean 255 mm vs. 285 mm). It has lower mean counts of ventral scales (76 vs. 85) than A. ceriacoi. The ranges of this character slightly overlap in the two species, although they are higher in A. ceriacoi sp. nov. than in A. cyanogaster. The mean number of precloacal pores is much higher in A. cyanogaster than in A. ceriacoi (38 vs 18). [A. cyanogaster is not a part of A. atricollis group but is included here due to its morphological similarity and because of the historical use of the name to describe this population].
(e) Acanthocercus gregorii has a smaller SVL (mean 119 mm vs. 123 mm) but greater total length (mean 291 mm vs. 285 mm). A. gregorii has lower mean counts of scale rows around midbody (104 vs. 108) than A. ceriacoi. The ranges of these characters are higher in A. ceriacoi than in A. gregorii. The mean number of precloacal pores is higher in A. gregorii than in A. ceriacoi (22 vs 18). In displaying adult males, A. gregorii are characterized by their bluish-green head and a uniform greenish-blue throat, with a uniform bluebody (Klausewitz 1957; Wagner et al. 2018), whereas in A. ceriacoi the blue of the male is limited to
the head.
(f) Acanthocercus kiwuensis has a smaller SVL (mean 111 mm vs. 123 mm) but greater total length (mean 290 mm vs. 285 mm). A. kiwuensis has lower mean counts of scale rows around midbody (87 vs. 108) and ventral scales (67 vs. 85) than A. ceriacoi. The ranges of this character largely overlap in the two species, although they are higher in A. ceriacoi than in A. kiwuensis. Moreover A. kiwuensis does not possess transverse rows of enlarged scales (Wagner et al. 2018). The mean number of precloacal pores is lower in A. kiwuensis than in A. ceriacoi (10 vs 18).
(g) Acanthocercus margaritae has a smaller SVL (mean 112 mm vs. 123 mm) and total length (mean 258 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (136 vs. 108), dorsal scales (103 vs. 63), and ventral scales (101 vs. 85) than A. ceriacoi sp. nov.. The ranges of these characters are lower in A. ceriacoi than in A. margaritae. The mean number of precloacal pores is lower in A. margaritae than in A. ceriacoi (14 vs 18).
(h) Acanthocercus minutus has a smaller SVL (mean 101 mm vs. 123 mm) and total length (mean 234 mm vs. 285 mm). It has higher mean counts of dorsal scales (79 vs. 63), and lower mean counts of ventral scales (78 vs. 85) than A. ceriacoi . The mean number of precloacal pores is higher in A. minutus than in A. ceriacoi (26 vs 18).
(i) Acanthocercus ugandaensis has a smaller SVL (mean 103 mm vs. 123 mm) and total length (mean 275 mm vs. 285 mm). A. ugandaensis has lower mean counts of scale rows around midbody (94 vs. 108) and ventral scales (73 vs. 85) than A. ceriacoi. The ranges of these characters are higher in A. ceriacoi than in A. ugandaensis. The mean number of precloacal pores is higher in A. ugandaensis than in A. ceriacoi (24 vs 18). (Marques et al. 2022)

Coloration. In displaying males head vibrantly blue; neck, shoulders, body and limbs brownish with a pattern of white and black stripes and speckles; a pale vertebral band from the neck to the tail is present, distinct anteriorly between the flanks, becoming more indistinct and on a brown background posteriorly; enlarged dorsal scales of the lower part of the body yellowish, matrix scales greyish becoming brownish on the hind limbs; first half of the tail yellowish, second half of the tail with dark and lighter blue bands. Ventral parts of the body, limbs and tail dirty whitish with a brown patterning coloration on females in life. Shoulders brownish with a black patch, some individuals present a pale blue head. Dark brown, broad transverse bands between limbs, each interspersed by transverse bands composed of a single row of enlarged yellow scales.Ventral parts of the body, limbs and tail dirty whitish with an extensive dark brown pattern. Gular region of a vibrant blue. (Marques et al. 2022) 
CommentDistribution: Likely to occur in the neighboring provinces of Kwanza-Norte and Zaire, as well as in southern Democratic Republic of the Congo.

Specimens of Acanthocercus have been recorded from Angola since the mid nineteenth century under a variety of names. Bocage (1866) referred to the species as Stellio nigricollis, a lapsus calami for atricollis, as noted later by Bocage (1895) and confirmed by Wagner et al. (2018). Subsequently, the species has been recorded as Stellio atricollis (Bocage 1879, 1895; Peters 1881; Boulenger 1885, 1905; Ferreira 1900; Angel 1923; Schmidt 1933; Parker 1936; Monard 1937; Themido 1941), Agama atricollis (Boulenger 1885, 1905; Laurent 1950, 1964; Ferreira 1903; Schmidt 1933; Monard 1937; Parker 1936; Themido 1941; Frade 1963; Manaças 1963), and Agama/Acanthocercus cyanogaster [sic] (Loveridge 1957; Branch & Conradie 2015). In a generalist publication, Falk (1925) described a new species based on an unspecified Angolan population—Agama cyanocephala. As Falk (1925) did not provide any specific type locality and the whereabouts of any type material are unknown, in order to stabilize the taxonomy and nomenclature of the group, Wagner et al. (2018) designated a neotype from Ikelenge [-11.241592°, 24.273256°, 1420 m a.s.l.], northwestern Zambia and considered that most Angolan populations would represent this taxon. Following this interpretation from Wagner et al. (2018), subsequent authors who have dealt with Angolan populations of Acanthocercus have opted to identify the specimens at their disposal as A. cyanocephalus (e.g., Ceríaco et al. 2014, 2016). After preliminary analysis of freshly collected material from southern Angola (Huíla Province), Marques et al. (2018) divided the Angolan populations of Acanthocercus in two putative taxa, A. cyanocephalus in the central and northern regions of the country, and A. sp. in the southern provinces of Huíla, Cunene and Cuando-Cubango. More recently Wagner et al. (2021) revisited the Angolan and northern Namibian populations of the genus. Based on a taxonomic integrative approach they concluded that three distinct lineages of Acanthocercus occur in Angola: A. cyanocephalus (northeastern Angola), A. margaritae, (southern Angola and neighboring northern Namibia), and a third, that correspond to A. ceriacoi (central and northwestern Angola). (Marques et al. 2022) 
EtymologyNamed after Portuguese zoologist Luis Miguel Pires Ceríaco (1987–present), head of collections and research at the Museu de História Natural e da Ciência da Universidade do Porto, Porto, Portugal, and invited curator of herpetology at the Museu Nacional de História Natural e da Ciência, Lisbon, Portugal. Luis M.P. Ceríaco has been a major force comprising the new wave of Angolan herpetological studies and a dedicated supervisor to a new generation of Portuguese and Angolan herpetologists. The name is formed in the genitive masculine singular. 
References
  • MARQUES, M. P., PARRINHA, D., SANTOS, B. S., BANDEIRA, S., BUTLER, B. O., SOUSA, A. C. A., ... & WAGNER, P. 2022. All in all it’s just another branch in the tree: A new species of Acanthocercus Fitzinger, 1843 (Squamata: Agamidae), from Angola. Zootaxa 5099 (2), 221-243 - get paper here
 
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