Cytosystematics of hyperoliid frogs: Phylogeny of Heterixalus, low karyotypic variability in hyperoliines and separate phylogenetic position of Leptopelis

Figures & data

Table I. Summary of previous knowledge on chromosome numbers of hyperoliid frogs. For detailed references, see King (1990). Semnodactylus weali was originally considered to belong to the genus Kassina and its karyotype described as Kassina weali.

	No. species studied	No. chromosomes (2n)	No. sp. applied banding
Hyperolius	16	24	–
Heterixalus	2	24	–
Opisthothylax	1	24	–
Tachynemis	1	24	–
Afrixalus	6	24	–
Kassina	2	24	–
Semnodactylus	1	24	1 (Ag‐NOR, CMA3)
Phlyctimantis	1	24	–
Leptopelis	11	22–24–30	1 (Ag‐NOR, CMA3)

Figure 1 Giemsa stained karyotypes of the 13 studied hyperoliids.A, Heterixalus alboguttatus; B, H. andrakata; C, H. betsileo; D, H. boettgeri; E, H. luteostriatus; F, H. punctatus; G, H. rutenbergi; H, H. tricolor; I, H. variabilis; J, Acanthixalus spinosus; K, Hyperolius cf. viridiflavus; L, Kassina maculata; M, Leptopelis calcaratus. The Ag‐NOR banded pairs are reported above the corresponding Giemsa stained pairs.

Table II. Relative lengths (±SD; upper value) and centromerix indices (±SD; lower value) of chromosomes 1–12 in hyperoliid karyotypes studied.

Chromosome	1	2	3	4	5	6	7	8	9	10	11	12
H. alboguttatus	14.7±0.5	11.8±0.7	11.4±0.8	10.5±0.4	10.5±0.6	9.1±0.7	6.8±0.5	6.3±0.6	5.3±0.5	4.9±0.7	4.7±0.7	4.6±0.8
	48.8±4.7	34.2±3.2	32.4±3.8	32.5±3.0	48.7±2.9	47.5±3.0	48.0±3.7	45.4±3.3	42.9±4.5	41.7±5.0	41.6±3.6	44.6±2.8
H. andrakata	16.5±0.6	10.6±0.8	10.6±0.4	10.3±0.7	10.3±0.5	7.0±0.6	6.5±0.6	6.5±0.5	6.4±0.7	5.7±0.8	5.2±0.6	4.4±0.6
	47.9±4.1	30.8±3.9	35.6±4.0	33.8±4.6	40.1±3.5	44.8±4.0	45.9±3.9	47.5±4.3	48.8±4.9	45.6±4.4	43.8±4.1	45.6±3.9
H. betsileo	14.3±0.5	11.5±0.6	11.4±0.5	9.8±0.6	9.8±0.7	7.4±0.9	6.9±0.5	6.6±0.7	6.1±0.9	6.0±0.6	5.2±0.7	5.0±0.6
	46.8±3.9	31.8±3.7	34.9±2.7	30.9±4.0	41.4±3.5	43.8±3.5	43.9±3.8	45.8±3.0	42.9±4.3	46.8±2.8	43.8±3.0	47.2±3.7
H. boettgeri	15.0±0.5	12.2±0.7	11.0±0.6	10.2±0.8	10.2±0.5	8.4±0.6	6.3±0.8	6.1±0.4	5.5±0.8	5.5±0.7	5.3±0.5	4.3±0.4
	43.2±4.6	30.8±3.1	34.9±3.6	32.7±3.2	45.6±4.3	44.7±3.6	44.0±3.9	43.8±4.3	46.9±4.4	47.3±3.7	38.8±4.0	45.3±3.3
H. luteostriatus	13.7±0.6	11.0±0.5	10.7±0.8	10.3±0.5	10.3±0.7	8.3±0.5	6.5±0.6	6.4±0.8	6.0±0.9	5.9±0.8	5.8±0.6	5.1±0.6
	41.9±3.9	39.9±4.0	31.8±3.7	32.9±3.5	44.2±2.0	43.8±2.9	46.5±3.2	39.9±3.1	47.2±3.9	44.2±3.5	40.1±2.9	46.3±3.5
H. punctatus	13.7±0.6	11.3±0.7	10.9±0.6	10.1±0.5	9.9±0.6	8.3±0.8	7.9±0.6	6.4±0.6	6.1±0.7	5.9±0.5	4.8±0.6	4.7±0.8
	44.0±3.0	40.4±4.2	30.0±3.5	34.2±2.9	40.3±3.2	44.8±3.0	45.3±2.6	43.9±3.3	45.0±3.0	47.5±2.9	39.7±2.7	46.8±3.1
H rutenbergi	14.4±0.7	12.0±0.8	11.4±0.6	9.5±0.7	9.3±0.8	6.9±0.6	6.8±0.7	6.3±0.5	6.2±0.4	5.9±0.6	5.7±0.5	5.6±0.4
	43.2±3.2	33.8±3.0	32.7±2.8	31.9±3.0	42.5±3.4	43.2±2.8	44.8±3.8	46.7±3.6	45.3±4.0	42.8±2.9	46.0±3.2	44.8±3.5
H. tricolor	13.6±0.5	12.7±0.6	12.1±0.7	9.7±0.6	9.1±0.5	8.4±0.8	7.6±0.6	6.5±0.5	5.2±0.7	5.8±0.9	5.0±0.6	4.3±0.5
	41.9±3.0	32.8±2.9	34.8±3.8	32.0±3.5	42.8±3.1	44.5±2.9	45.7±2.2	46.3±3.8	43.8±4.0	48.4±2.9	43.2±3.0	44.8±2.9
H. variabilis	13.1±0.4	10.4±0.4	10.1±0.7	9.6±0.6	9.4±0.7	9.1±0.5	7.8±0.7	7.2±0.8	6.5±0.6	6.2±0.7	5.8±0.6	5.0±0.6
	43.8±3.8	32.5±3.0	32.0±2.7	31.5±3.0	44.3±2.7	45.8±3.3	46.9±3.2	47.6±2.9	43.8±4.1	44.9±2.9	45.0±3.5	46.8±2.5
A. spinosus	15.3±0.6	11.5±0.6	10.4±0.5	10.0±0.6	9.3±0.5	8.6±0.7	8.0±0.6	6.1±0.6	5.9±0.8	5.8±0.4	5.1±0.6	4.0±0.5
	40.0±3.2	35.8±3.0	33.8±2.9	34.9±2.7	45.3±3.3	45.2±2.8	42.8±3.2	44.7±3.8	45.3±2.7	40.6±3.0	43.5±4.1	38.9±3.5
Hy. cf. viridiflavus	15.0±0.5	10.9±0.8	10.8±0.7	9.7±0.6	9.5±0.6	8.0±0.7	6.9±0.5	6.7±0.8	6.3±0.7	5.8±0.6	5.4±0.7	5.0±0.5
	43.4±3.3	32.0±3.2	29.4±2.8	31.2±3.3	41.1±2.8	42.6±3.0	42.2±2.0	44.4±3.5	46.8±3.4	41.9±3.7	43.3±2.9	45.0±3.0
K. maculata	13.0±0.6	11.5±0.7	10.4±0.6	9.9±0.6	9.9±0.5	7.9±0.4	7.8±0.6	6.3±0.7	6.3±0.6	6.2±0.5	6.1±0.4	4.7±0.8
	45.2±2.8	30.2±2.8	30.9±3.0	32.1±3.1	43.6±2.9	43.4±2.9	46.8±2.7	48.6±3.9	41.9±3.6	38.0±3.2	46.3±3.	24.2±2.8
L. calcaratus	15.5±0.7	11.5±0.5	11.0±0.6	10.4±0.5	10.3±0.7	7.7±0.6	6.4 ±0.8	6.0±0.5	5.7±0.6	6.1±0.4	6.0±0.8	3.4±0.7
	45.3±2.7	26.9±3.8	29.0±3.4	36.0±2.8	43.2±3.8	44.9±2.	43.3±2.0	43.5±3.8	42.9±3.9	36.8±2.5	46.3±3.6	0.0

Figure 2 C‐banded karyotypes of the 13 studied studied hyperoliids. A, Heterixalus alboguttatus; B, H. andrakata; C, H. betsileo D, H. boettgeri; E, H. luteostriatus; F, H. punctatus; G, H. rutenbergi; H, H. tricolor; I, H. variabilis; J, Acanthixalus spinosus; K, Hyperolius cf. viridiflavus; L, Kassina maculata; M, Leptopelis calcaratus.

Figure 3 Sequential C‐banding (rows I and II) or Alu I (rows III and IV)+CMA₃ (rows I and III)+DAPI (rows II and IV) of: A, H. alboguttatus; B, H. andrakata; C, H. betsileo; D, H. boettgeri; E, H. luteostriatus; and F, H. punctatus. The colour version of this figure is available online.

Figure 4 Sequential C‐banding ( rows I and II) or Alu I (rows III and IV)+CMA₃ (rows I and III)+DAPI (rows II and IV) of A, H. rutenbergi; B, H. tricolor; C, H. variabilis; D, Acanthixalus spinosus; E, Hyperolius cf. viridiflavus; F, Kassina maculata; and G, Leptopelis calcaratus. The colour version of this figure is available online.

Table III. Summary of karyotype data resulting from the present study. Abbreviations used: Cen = centromeric; Tel = Telomeric; Pcen = Paracentromeric; m = metacentric; sm = submetacentric; p→int = NORs interstitial on short arm; q→stel = NORs peritelomeric on long arm; p→tel = NORs telomeric on short arm; q→tel = NORs telomeric on long arm; p→cen = pericentromeric on long arm; Neg = negative; pr/prs = pair(s).

Species	2n	2nd pair	NORs	C‐banding+Giemsa	Paracentromeric C‐bands	C‐banding+CMA3+DAPI	Alu I‐banding+CMA3+DAPI	NOR associated heterocromatin
Heterixalus
H. alboguttatus	24	sm	9th; q→stel	Cen (+5 prs), Tel (+4 prs)	Pairs 2,3,5,6,7,8	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
H. andrakata	24	sm	9th; q→stel	Cen (+all prs), Tel±	Pairs 5 and 8	Neg +(3 prs)	Cen−, T+Neg	C±, CMA3+, Alu I+
H. betsileo	24	sm	9th; q→stel	Cen (+8 prs), Tel (+2 prs)	None	Cen (+2 prs) Neg	Neg Neg	C±, CMA3+, Alu I+
H. boettgeri	24	sm	9th; q→stel	Cen (+1 pr, Tel (+10 prs)	Pairs 3,5,6, 8	Pcen+Pcen+	Pcen+Pcen+	C+, CMA3+, Alu I+
H. luteostriatus	24	m	9th; p→int	Cen (+2 prs), Tel (+all prs)	None	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
H. punctatus	24	m	9th; p→int	Cen (+9 prs), Tel−	None	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
H. rutenbergi	24	sm	9th; p→int	Cen (+all pairs), Tel−	None	Neg Neg	Neg Neg	C±, CMA3+, Alu I+
H. tricolor	24	sm	9th; q→stel	Cen (+all pairs), Tel (+2 prs)	None	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
H. variabilis	24	sm	9th; q→stel	Cen (+4 prs), Tel (+10 prs)	None	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
Acanthixalus
A. spinosus	24	sm	9th; p→tel	Cen (+all prs), Tel (++5th pr)	None	Neg Neg	Neg Neg	C−, CMA3−, Alu I−
Hyperolius
H. cf. viridiflavus	24	sm	9th; q→tel	Cen (+2prs), Tel (+7 prs)	None	Neg Neg	Neg Neg	C+, CMA3+, Alu I+
Kassina
K. maculata	24	sm	9th; q→tel	Cen (+all prs), Tel (+3 prs)	None	Cen+Cen+	Neg Neg	C−, CMA3−, Alu I−
Leptopelis
L. calcaratus	24	sm	5th; p→cen	Cen (+all prs),Tel (+all prs)	Pair 1	Neg Neg	Cen+ Cen+	C−, CMA3−, Alu I−

King , M. 1990 . “ Amphibia, Vol. 4 Chordata 2. ” . In Animal Cytogenetics. Stuttgart , Edited by: John , B . 1 – 241 . Berlin : Gebrüder Borntraeger .

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