Corresponding author: Stuart V. Nielsen (
Academic editor: A. Crottini
Recent molecular phylogenetic work has found that
Investigações moleculares recentes revelaram que o género
Nielsen SV, Conradie W, Ceríaco LMP, Bauer AM, Heinicke MP, Stanley EL, Blackburn DC (2020) A new species of Rain Frog (Brevicipitidae,
The taxonomy of Angolan
A recent phylogenetic study of
Angola’s long civil war, which lasted from 1975 to 2002, effectively stifled biological exploration and discovery (for additional summary, see
We consider species as units of separately evolving metapopulation lineages, following the conceptual framework developed by
Between 2016 and 2019, specimens referable to the genus
Sampling information including specimens/field IDs (Museum Abbreviations:
species | Tree ID | Specimen ID | Field ID | Latitude and Longitude | Country | Locality |
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12S | 16S |
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ANG-01 | UF Herp 187172 |
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ANG | Kawa Camp (1 km S of the Kwanza R.), Kissama NP, Luanda Prov. |
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ANG-02 | UF Herp 187173 |
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ANG | Kawa Camp (1 km S of the Kwanza R.), Kissama NP, Luanda Prov. |
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ANG-03 | MHNCUP_ANF 0320 | AMB11736 |
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ANG | Embala Seque (14 km N of Cassumbi village), Bie Province |
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ANG-04 | WC-3924 |
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ANG | Cuito River source lake, Moxico Province |
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ANG-05 | WC-4591 |
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ANG | Cuanavale River source lake, Moxico Province |
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ANG-06 | WC-4756 |
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ANG | Quembo River source lake, Moxico Province |
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ANG-07 | WC-4827 |
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ANG | Cuando River source, Moxico Province | – | – |
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ADS-01 | AMB8318 |
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RSA | Farm Celine, Limpopo |
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– |
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ADS-02 |
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ZIM | Hwange |
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– |
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ADS-03 |
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ZIM | Miombo Safari Camp |
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– |
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ADS-04 |
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NAM | Farm Ohange, Otjozondjupa | – | – |
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ADS-05 | – | SVN 766 |
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RSA | Ellisras |
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ADS-06 | – | SVN 768 |
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RSA | Ellisras |
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ADS-07 | AMB7963 |
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NAM | Katima Mulilo |
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ADS-08 | – | AMB7972 |
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NAM | Caprivi |
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ADS-09 | AMB7980 |
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NAM | Caprivi |
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MOS-01 | DMP 344 |
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MOZ | Gurue |
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MOS-02 |
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Mulanje |
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MOS-03 | RB09-159 |
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MOZ | Ila de Mozambique |
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MOS-04 | RB09-179 |
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MOZ | Ila de Mozambique |
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MOS-05 | RB09-030 |
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MOZ | Pemba |
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MOS-06 | RB09-046 |
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MOZ | Pemba |
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MOS-07 | RB10-A097 |
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MOZ | Nampula |
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MOS-08 | NIMB 112 |
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MOZ | Lichinga |
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MOS-09 | Syran 12 |
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MOZ | Balama |
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POW-01 | – | ELI 325 |
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DRC | Manono |
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POW-02 | JWH10-A114 |
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ZAM | Kalumbila |
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POW-03 | RB10-F003 |
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ZAM | Lusaka |
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POW-04 | RB10-F012 |
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ZAM | Lusaka |
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We amplified partial sequences of two mitochondrial (12S and 16S ribosomal rRNA genes) and two nuclear loci (recombination activating protein 1,
Uncorrected mean pairwise 12S and 16S mitochondrial sequence differences between ingroup
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0.09 | 0.09 | 0.11 |
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0.11 |
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0.09 | 0.08 | |
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0.12 | 0.11 |
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0.09 | |
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0.12 | 0.08 | 0.10 |
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Datasets (concatenated and partitioned by locus/codon) of all samples were analyzed using maximum likelihood (RAxML v.8.2;
Specimens were measured to the nearest 0.1 mm using digital calipers under a dissecting stereomicroscope for the following 24 morphological characters as defined by
Advertisement calls were recorded in the field using an Samsung Galaxy Note 3 cellphone at a sampling rate of 44100
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature (
Our concatenated, multi-locus dataset was 1,852 bp long, of which 390 characters were parsimony informative. Phylogenetic analyses resulted in a well-supported species-level phylogeny and high support that
Geographic distribution and phylogenetic relationships of
Mensural and meristic data are presented in Table
Measurements (mm) of type series.
UF Herp 187172 | UF Herp 187173 | MHNCUP_ANF 0320 | Average | SD | ||||||
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30.5 | 27.5 | 24.6 | 25.4 | 18.3 | 26.6 | 30.1 | 26.5 |
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26.3 | 24.9 | – | 23 | 17.6 | 23.2 | 29 | 25.2 |
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7.2 | 7.3 | 7.3 | 10.8 | 6.4 | 9.6 | 12.7 | 9.3 |
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3.2 | 2.8 | 2.9 | 3.1 | 2.2 | 2.7 | 3.3 | 2.5 |
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2.0 | 1.7 | 1.6 | 2.1 | 2.5 | 1.7 | 2.2 | 2.1 |
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2.7 | 2.8 | 3.2 | 3.4 | 1.3 | 3.1 | 3.6 | 2.8 |
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1.7 | 1.7 | 1.3 | 1.4 | 1 | 1.2 | 1.8 | 1.3 |
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1.7 | 1.3 | 1.7 | 1.8 | 1.4 | 2.1 | 2.3 | 1.9 |
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1.9 | 1.9 | 2 | 1.6 | 1.4 | 1.8 | 2.2 | 2 |
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7.2 | 6.8 | 5.1 | 6.9 | 4.7 | 6.3 | 7.9 | 7 |
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EAD | 9.1 | 9.3 | 4.1 | 4.5 | 3.6 | 4.3 | 5.3 | – |
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1.7 | 1.5 | 1.5 | 2.7 | 1.7 | 2.7 | 2.5 | 2.8 |
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1.9 | 1.9 | 1.7 | 3.6 | 1.8 | 3.2 | 2.6 | 2.7 |
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2.9 | 3.0 | 2.2 | 3.9 | 2.6 | 3.9 | 3.6 | 3.7 |
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1.2 | 1.3 | 1.2 | 1.8 | 0.9 | 2 | 1.5 | 1.4 |
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1.0 | 1.1 | 1 | 1.4 | 0.6 | 1.5 | 1.6 | 1.2 |
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1.8 | 1.9 | 2.9 | 2.4 | 2 | 2.7 | 1.5 | 2.7 |
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4.4 | 4.2 | 4.6 | 4.1 | 3.2 | 5 | 4.9 | 4.6 |
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FT | 10.9 | 10.4 | 8 | 8.9 | 6 | 10.3 | 10.5 | 9.7 |
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4.0 | 5.0 | 0.8 | 4 | 3.2 | 4.9 | 5.3 | 4.7 |
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MTL | 1.0 | 1.0 | 1.2 | 2.3 | 1.7 | 2.3 | – | – |
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3.0 | 2.9 | 3 | 3.2 | 2.3 | 3.4 | 3.6 | 3.7 |
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TIB | 8.5 | 8.3 | – | 6.4 | 4.8 | 7.4 | 8.8 | 8.1 |
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The advertisement call of the eastern population is pulsed, has a call duration of 0.175 ± 0.083 s, with relatively long intervals between consecutive calls (0.996 ± 0.133 s), a high number of pulses per call (28–34; Table
Spectrograms and oscillograms showing a series of notes of the putatively novel Angolan
Principal components analysis (
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Proportion of Variance | 29.92 | 26.57 | 13.70 | 8.67 | |
Cumulative Proportion | 29.92 | 56.49 | 70.19 | 78.86 | |
Loadings | |||||
Head Length ( |
-0.3025245 | 0.35446298 | -0.3789378 | 0.02808779 | W = 0.943, p = 0.083 |
Head Width ( |
0.33281314 | 0.38971121 | 0.08705045 | 0.33211061 | W = 0.956, p = 0.205 |
Eye diameter ( |
-0.2210569 | 0.22060777 | -0.6170786 | -0.1165699 | W = 0.963, p = 0.312 |
Snout length ( |
-0.4530121 | -0.0316756 | 0.28394189 | -0.1665634 | W = 0.973, p = 0.567 |
Interorbital distance ( |
-0.3043034 | 0.24152256 | 0.52046607 | 0.21274977 | W = 0.959, p = 0.240 |
Internarial distance ( |
-0.4018886 | 0.1543163 | 0.09759718 | 0.52889612 | W = 0.965, p = 0.360 |
Thigh length ( |
0.2280386 | 0.3347664 | 0.24257868 | -0.4343642 | W = 0.965, p = 0.360 |
Crus length ( |
0.45829394 | 0.12745601 | 0.05499745 | 0.20448849 | W = 0.975, p = 0.636 |
Pes length (FT) | -0.1152479 | 0.43230304 | 0.17838021 | -0.5130485 | W = 0.900, p = 0.005 |
Manual digit III length ( |
0.11914774 | 0.52484652 | -0.115415 | 0.16994577 | W = 0.989, p = 0.978 |
Our phylogenetic analyses indicate that sampled individuals from Angola form a clade that is genealogically exclusive from other described species of
A species referable to
The new species can be distinguished from other species of
The advertisement call of the new species (Table
Comparison of the main variables for the advertisement calls of
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avg ± sd | range | avg ± sd | range | avg ± sd | range | avg ± sd | range | |
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0.175 ± 0.083 | 0.064–0.342 | 0.196 ± 0.047 | 0.077–0.293 | 0.500 ± 0.070 | 0.036–0.079 | 0.140 ± 0.012 | 0.111–0.160 |
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0.996 ± 0.133 | 0.742–1.190 | 0.745 ± 0.636 | 0.363–0.745* | 0.710 ± 0.168 | 0.396–1.17 | 0.743 ± 0.166 | 0.500–1.100 |
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30 ± 2.6 | 28–34 | 23 ± 3.3 | 14–31 | 9 ± 1.2 | 7–13 | 30 ± 16.3 | 10–74 |
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2156 | na | 1742 ± 100 | 1482–2179 | 1835 ± 107 | 1600–2193 | 1728 ± 83 | 1557–1903 |
Adult male (
Skin of dorsum and head smooth, and weakly glandular with irregular folds; skin of ventrum smooth; skin folds overlying vent creating triangular shape.
Limbs short with digits I and V short or rudimentary; webbing absent on manus and pes; nuptial pads absent and adhesive glands not discernable; relative manual digit lengths when adpressed: III>II>I>IV; only tip of first pedal digit extending beyond fleshy webbing and sole; fourth (outer) manual digit reaches midway between the large tubercle at metacarpophalangeal joint and subarticular tubercle at most proximal interphalangeal joint; finger tips conical, not expanded; several small globular palmar tubercles; single subarticular tubercles present on pedal digits II, III, and IV; pedal digit V very short, falling short of most proximal subarticular tubercle of pedal digit IV; well-developed (though not keratinized) inner metatarsal tubercle visibly longer than pedal digit III, separated from conical outer metatarsal tubercle by deep cleft.
In life, dorsum of body mottled dark brown on pale tan base, transitioning to golden yellow on the lateral aspects, before stark transition to solid dark brown flanks with a dark boundary becoming paler ventrally (Fig.
In preservative, coloration is largely similar but more muted and overall darker (Fig.
Measurements of the type series are shown in Table
All specimens resemble the holotype in the absence of a visible tympanum, and skin that is densely granular dorsally and laterally and smooth ventrally (Figs
Variation in color and pattern within living paratypes of
Color and pattern in UF Herp 187173 is very similar to the holotype. Dorsum gray with scattered black spots (
Variation in color and pattern within preserved paratypes of
The following call description is based on a recording of a paratype male (
Based on our phylogenetic analysis, this species is currently confirmed from three widely separated localities and elevations ranging from near sea level to > 1400 m: i) Kissama National Park, on the outskirts of Angola’s capital city, Luanda, in coastal western Angola (Luanda Province); ii) central Angola (Bié Province); and iii) the source of the Cuanavale, Cuito, Cuando and Quembo rivers (Moxico Province). The identity of other known Angolan localities for
The preferred habitat for
Photos of typical habitat of
The name
Given that it appears widely distributed, we suggest that
Further work is required to confirm the distributional range of
We are not the first to recognize the lack of morphological variation within members of this anuran clade, which has led to historical taxonomic confusion and invoking hybridization for specimens that failed to conform to often scant descriptions of the type specimens (
As mentioned above, there is considerable genetic structure within
We are grateful to Mariana P. Marques, Adam Ferguson, Ben Marks, John Cavagnaro, Philip Pastor, Suzana Bandeira, Ilola Jorge, Alvaro “Varito” Baptista, Ninda Baptista, Kerllen Costa, James Harvey, Roger Bills, Götz Neef, and Luke Verburgt for their invaluable field assistance, support, advice, and companionship. John Cavagnaro provided the life photo of the holotype. Mohamad Beidoun assisted with molecular work. This work was supported in part by the US National Science Foundation (DEB-1556255, 1556559 and DEB-1556585 to DCB, AMB, and MPH), the JRS Biodiversity Foundation (to DCB and AMB), and the National Geographic Society (Okavango Wilderness Project EC0715-15 to WC). The funders had no role in study design, data collection, and analysis, the decision to publish, or manuscript preparation. Field components of this work were facilitated by a Memorandum of Understanding between the University of Florida and The National Institute for Biodiversity and Protected Areas (INBAC) in Angola. Material used in this study was exported under the following permit numbers: 083/INBAC.MINAMB/2016 (to Villanova University) and 31/GGPCC/2016 (to the
We here provide only the usages that have been applied to Angola populations, albeit with some inherent uncertainty given the pervasive morphological similarity among members of the B. mossambicus group.
Table S1. Morphological data used to perform PCAs
morphological data
Morphological data used to perform PCAs. See Table