hylogenetic relationships and phylogeography of the KilliWsh
species of the subgenus Chromaphyosemion (Radda, 1971)
in West Africa, inferred from mitochondrial DNA sequences
J.-F. Agnèse a,b,¤, F. Zentz c, O. Legros d, D. Sellos e
a Laboratoire Génome, Populations, Interactions, Adaptation, UM2, IFREMER-CNRS UMR5000, Université de Montpellier II,
cc63, place E. Bataillon, F34095 Montpellier Cedex 5, France
b Institut de Recherche pour le Développement, 213 rue La Fayettte, 75480 Paris Cedex 10, France
c Université de Bretagne Occidentale, IUT de Quimper, Dpt Génie Biologique, 2 rue de l’Université, 29334 Quimper Cedex, France
d Floralaan, 51, 1501 Buizingen, Belgium
e Station de Biologie Marine de Concarneau, Muséum National d’Histoire Naturelle, UMR 5178, BP 225, 29182 Concarneau Cedex, France
Received 10 March 2006; accepted 14 March 2006
Available online 19 April 2006
Fig. 1. Map of West Africa, showing approximate distribution ranges of the 12 species of the subgenus Chromaphyosemion (grey zone in the small map)
with sampling locations. White circles, locations for wild samples collected for this study; black circles, locations for aquarium strain specimens. Even if
species ranges overlap, the diVerent species are not sympatric except in Wve locations (not shown on the map). The isolated population of A. aV splendopleure
Ekom falls (not studied here) is indicated by a star. The question mark indicates a zone where Chromaphyosemion are likely to be present but where
Wsh have never been collected because of the absence of road. The hypothesized refuge zone was indicated in black.
Table 1
Population number (No), origin, locality, coordinates, species identiWcation, previous determination (PrDet), and voucher numbers
No Origin Locality Coordinates Species Pr.Det Voucher number
1 AS Ibadan, 47 km of Lagos 7.400°N, 3.900°E BIT
2 AS Umudike 5.310°N, 7.250°E BIT
3 AS Kwa River Falls Plantation 5.070°N, 8.200°E BIV
4 AS Biafra 4.750°N, 8.920°E BIV
5 AS Funge 4.764°N, 8.907°E BIV
6 ABC 05/12 Funge 4.764°N, 8.907°E BIV 2005-16-P-19-20
7 ABC 05/13 Ekondo Nene 4.690°N, 8.966°E VOL SPP 2005-16-P-21-22
8 ABC 05/14 Kumba 4.654°N, 9.506°E VOL SPP 2005-16-P-23-24
9 ABC 05/15 Two Wata 4.694°N, 9.791°E VOL SPP 2005-16-P-25-26
10 ABC 05/16 Sole 4.602°N, 9.814°E VOL SPP 2005-16-P-27-28
11 AS Lykoko 4.400°N, 9.330°E VOL SPP
12 ABC 05/10 Muyuka 4.380°N, 9.439°E VOL SPP 2005-16-P-17-18
13 ABC 05/08 Mile 29 4.250°N, 9.360°E VOL PLKaV 2005-16-P-15-16
14 AS Ekona CBL 01/22 4.340°N, 9.561°E PLK
15 ABC 05/07 3 km North Buea cross road 4.254°N, 9.207°E PLK 2005-16-P-13-14
16 AS Monea 4.200°N, 9.300°E PLK
17 AS Bolifamba 4.150°N, 9.300°E PLK
18 AS CBL 01/23 4.138°N, 9.395°E PLK
19 ABC 05/06 North of Moliwe 4.182°N, 9.308°E VOL SPP 2005-16-P-11-12
20 AS Rio Consul GEMHS 00/43 3.993°N, 9.053°E SPP SPBK
21 AS Bimbia Camp 3.854°N, 9.184°E SPP SPBC
22 AS Entrance to Tiko CBL 01/25 4.298°N, 9.474°E SPP
23 ABC 05/05 7.5 km East of Tiko 4.185°N, 9.469°E VOL SPP 2005-16-P-9-10
24 AS Nkwo 97/1 4.516°N, 9.966°E RIG
25 ABC 05/18 13 km South Edea-Yabassi cross road 4.373°N, 10.018°E RIG 2005-16-P-29-30
26 ABC 05/19 Nkouli-Ngnock 4.183°N, 10.050°E RIG 2005-16-P-31
27 AS Bonepoupa-Yabassi km 20 C 89/18 4.371°N, 10.195°E RIG
28 ABC 05/01 North of Pouma 3.895°N, 10.556°E LOE 2005-16-P-1-2
29 ABC 05/02 17 km West of Pouma 3.823°N, 10.380°E LOE 2005-16-P-3-4
30 ABC 05/68 Song Ndong 3.823°N, 10.244°E LOE 2005-16-P-102-103
31 ABC 05/67 West of Song Ndong 3.800°N, 10.216°E LOE 2005-16-P-100-101
32 ABC 05/66 East of Batombe 3.809°N, 10.201°E LOE 2005-16-P-99
33 ABC 05/65 Batombe 3.826°N, 10.168°E LOE 2005-16-P-97-98
34 ABC 05/03 Song Ndong 3.673°N, 10.257°E LOE 2005-16-P-5-6
35 AS Edea-Yaounde km 18 C 89/31 3.490°N, 10.150°E LOE
36 AS Edea-Yaounde km 18 CSK 95/28 3.816°N, 10.246°E LOE
37 AS Nkakanzok CBL 01/13 3.958°N, 10.452°E LOE
38 ABC 05/64 East of Edea 3.773°N, 10.163°E LOE 2005-16-P-96
39 ABC 05/04 North of Edea 3.840°N, 10.087°E SPP 2005-16-P-7-8
40 ABC 05/21 Songueland 3.789°N, 10.071°E SPP 2005-16-P-33-34
41 AS Sipe CBL 01/15 3.795°N, 10.213°E SPP SPK
42 ABC 05/63 Koukoue 3.962°N, 10.104°E SPP SPK 2005-16-P-94-95
43 AS Apou C 89/30 3.380°N, 10.070°E LOE
44 ABC 05/20 Ndorbe River 3.786°N, 9.908°E SPP 2005-16-P-32
45 AS Ndog Bong CBL 01/10 4.007°N, 10.000°E SPP
46 AS Benengue CBL 01/11 3.819°N, 10.017°E SPP
47 ABC 05/62 North of Bonguen 3.525°N, 10.110°E SPP 2005-16-P-92-93
48 ABC 05/61 Bivouba 3.349°N, 10.098°E LOE 2005-16-P-90-91
49 ABC 05/60 Ebea 3.182°N, 10.028°E LOE 2005-16-P-88-89
50 ABC 05/59 Bipaga I 3.108°N, 9.991°E SPP
51 ABC 05/70 Mpolgue 3.030°N, 9.972°E LOE
52 ABC 05/38 Makouré II 3.048°N, 10.164°E LOE 2005-16-P-60-61
53 ABC 05/32 Bissiang 2.988°N, 10.000°E SPP + LOE 2005-16-P-51-52
54 ABC 05/33 Bissiang 2.986°N, 9.995°E LOE 2005-16-P-53-54
55 ABC 05/34 East of Bissiang 2.933°N, 10.012°E LOE 2005-16-P-55
56 ABC 05/35 Bidou I 3.044°N, 10.113°E LOE 2005-16-P-56-57
57 ABC 05/36 Bifoum 3.074°N, 10.389°E LOE PAP? 2005-16-P-58-59
58 AS Bidjouka BLLMC05/25 3.139°N, 10.595°E LOE PAP?
59 ABC 05/31 Akom II 2.813°N, 10.568°E LOE 2005-16-P-49-50
60 ABC 05/30 Assok I 2.887°N, 10.458°E LOE 2005-16-P-47-48
61 ABC 05/29 Aso I 2.789°N, 10.453°E LOE 2005-16-P-45-46
62 ABC 05/28 Fenda 2.817°N, 10.381°E LUGcf SP6 2005-16-P-43-44
63 ABC 05/27 Elon 2.807°N, 10.234°E LOE 2005-16-P-41-42
Table 1 (continued)
AS, aquarium strain; ABC 05/.., Wsh collected during a Wled trip in January 2005 by J.F. Agnèse, R. Brummett, and P. Caminade, the last number refer to
the station number during this collection trip; identically CBL 01/.., C89/.., GEMHS 2000/.., CSK 95/.., BLLMC 05/.., HLM 99/.., and LEC 93/.. refer to
other Weld trips. Details concerning all these collections can be found at http://www.killiWsh.f9.co.uk/KilliWsh/KilliWsh%20Website/Code_Library/
Code_Index.htm, or at http://www.chromaphyosemion.com or at http://chromaphyosemion.killi.org/foreigners/index.html. ALP, A. alpha; BIT, A. bitaeniatum;
BIV, A. bivittatum; KOU, A. kouamense; LUG, A. lugens; LUGcf, A. cf lugens; RIG, A. riggenbachi; LOE, A. loennbergii; MEL, A. melanogaster;
PAP, A. pappenheimi; PLK, A. poliaki; PLKaV, A. aV poliaki; SPP, A. splendopleure; SPBK, A. sp. Bioko; SPBC, A. sp. Bimbia Camp; SPK, A. sp.
Koukoue; SPL, A. sp. Likado; SPM, A. sp. Mboro; SP6, A. sp. No6; SPRM, undescribed species from Rio Muni; VOL, A. volcanum. All voucher numbers
refer to Wsh deposited at the Royal Museum of Central Africa, Tervuren, Belgium.
No Origin Locality Coordinates Species Pr.Det Voucher number
64 ABC 05/26 Adjap I 2.816°N, 10.164°E LOE 2005-16-P-39-40
65 ABC 05/25 Afanouan 2.833°N, 10.095°E LOE 2005-16-P-37-38
66 ABC 05/24 Nko olong 2.853°N, 10.033°E MEL 2005-16-P-35-36
67 ABC 05/23 West of Naha 2.885°N, 9.970°E MEL 2005-11-P-06-07
68 ABC 05/22 West of Nagadjo 2.900°N, 9.951°E MEL 2005-11-P-01-05
69 ABC 05/39 Ebome 2.899°N, 9.907°E MEL 2005-16-P-62-63
70 ABC 05/42 Mbode 2.749°N, 9.877°E SPP SPM 2005-16-P-64-65
71 ABC 05/43 North of Mboro 2.734°N, 9.878°E SPP SPM 2005-16-P-66-67
72 ABC 05/58 700m North of Lolabe River 2.687°N, 9.857°E MEL 2005-16-P-86-87
73 ABC 05/46 Near Ebodje 2.582°N, 9.848°E SPP SPL 2005-16-P-69-70
74 ABC 05/57 Mamelles region 2.492°N, 9.907°E SPP SPL 2005-16-P-85
75 ABC 05/47 Malaba 2.497°N, 9.836°E SPP SPL 2005-16-P-71
76 ABC 05/48 Afan Essokie II 2.464°N, 9.841°E SPP SPL 2005-16-P-72-73
77 ABC 05/49 Etonde Fang 2.427°N, 9.849°E PUN 2005-16-P-74-75
78 ABC 05/50 Pondo road (Campo) 2.371°N, 9.855°E PUN 2005-16-P-76-77
79 ABC 05/51 Bibabimvoto 2.362°N, 9.912°E PUN 2005-11-P-23-25
80 ABC 05/56 Akak 2.376°N, 9.964°E PUN 2005-16-P-83-84
81 ABC 05/52 Afan Essokie 2.381°N, 9.979°E LUG 2005-16-P-105-106
82 AS Afan Essokie HLM 99/28 2.395°N, 9.992°E LUG
83 ABC 05/53 East of Afan Essokie 2.382°N, 9.992°E LUG 2005-16-P-78
84 ABC 05/54 N’koleon 2.396°N, 10.043°E LUG 2005-16-P-79-80
85 ABC 05/55 Gate of the Campo Parc 2.372°N, 10.104°E LUG 2005-16-P-81-82
86 AS North Ndyiacom GEMHS 00/32 2.149°N, 10.005°E SPRM
87 AS 27.8 km NE Kougouleu LEC 93/24 0.250°N, 10.040°E KOU
88 AS 17,1 km to Cap Esterias LEC 93/26 0.566°N, 9.383°E ALP
Table 2
Pairwise divergence values (Kimura 2 genetic distances) between all species studied
DEL WAL EXU BUA CYA LAB MIM MAC CAM OCC AUR CIT COE LAM CHR COG DEC ELE MLP STR GAB
WAL .0883
EXU .1452 .1369
BUA .1375 .1372 .0217
CYA .1383 .1420 .0927 .0821
LAB .1297 .1364 .0883 .0988 .0989
MIM .1446 .1477 .1023 .1131 .1066 .0413
MAC .1487 .1477 .0990 .1133 .1033 .0381 .0187
CAM .1452 .1443 .1027 .1030 .1071 .0317 .0250 .0124
OCC .1456 .1450 .1131 .1063 .1099 .0675 .0811 .0745 .0610
AUR .1611 .1490 .1098 .1135 .1101 .0476 .0711 .0579 .0545 .0250
CIT .1526 .1482 .1207 .1245 .1134 .0642 .0781 .0714 .0646 .0443 .0347
COE .1301 .1296 .0890 .0962 .0893 .0446 .0480 .0416 .0447 .0541 .0379 .0543
LAM .1370 .1328 .0713 .0919 .0920 .0281 .0577 .0479 .0513 .0809 .0577 .0743 .0446
CHR .1370 .1401 .0781 .0919 .0920 .0313 .0610 .0578 .0546 .0843 .0677 .0776 .0578 .0155
COG .1367 .1361 .0816 .1025 .0957 .0314 .0611 .0513 .0547 .0845 .0611 .0778 .0513 .0093 .0124
DEC .1367 .1435 .0885 .1025 .0957 .0314 .0611 .0579 .0547 .0777 .0611 .0778 .0579 .0155 .0124 .0062
ELE .1330 .1398 .0850 .0990 .0957 .0282 .0578 .0546 .0514 .0811 .0644 .0744 .0546 .0124 .0093 .0031 .0031
MLP .1407 .1438 .0781 .0919 .0955 .0281 .0577 .0545 .0513 .0809 .0643 .0743 .0545 .0124 .0093 .0093 .0093 .0062
STR .1381 .1487 .1176 .1215 .1068 .0923 .0925 .0892 .0964 .0997 .0964 .1032 .0823 .0714 .0782 .0784 .0784 .0818 .0816
GAB .1542 .1649 .1215 .1253 .1183 .0929 .1072 .1039 .0966 .1034 .1072 .1034 .1034 .0719 .0719 .0789 .0720 .0754 .0753 .0251
OGO .1446 .1329 .0885 .0956 .0788 .0917 .1063 .1030 .1067 .1207 .0993 .1135 .0925 .0852 .0921 .0923 .0923 .0958 .0956 .0917 .0954
LOU .1483 .1293 .0881 .0986 .0751 .0883 .1093 .0990 .1027 .1166 .0993 .1025 .0995 .0781 .0815 .0816 .0816 .0850 .0849 .0952 .0954
HER .1603 .1634 .1251 .1253 .1109 .1176 .1251 .1253 .1220 .1217 .1256 .1145 .1145 .1176 .1212 .1251 .1251 .1251 .1248 .0956 .1067
EXD .1180 .1325 .0850 .0956 .0822 .0781 .0923 .0890 .0927 .1030 .0997 .0991 .0753 .0714 .0781 .0783 .0851 .0817 .0815 .0713 .0748
PRI .1177 .1249 .0921 .0958 .0855 .0848 .0921 .0888 .0925 .0993 .0960 .0989 .0717 .0780 .0848 .0849 .0918 .0884 .0882 .0611 .0818
ALH .1120 .1009 .0960 .0927 .0959 .0645 .0890 .0823 .0790 .1032 .0925 .1064 .0792 .0715 .0783 .0819 .0853 .0819 .0817 .0854 .0964
CAL .1161 .1156 .1069 .1179 .0998 .0647 .0964 .0896 .0863 .1032 .0925 .1067 .0896 .0785 .0785 .0750 .0716 .0750 .0820 .1036 .1079
AUS .1193 .1188 .0995 .0927 .0920 .0709 .0886 .0820 .0787 .0923 .0886 .1024 .0789 .0712 .0780 .0815 .0849 .0815 .0814 .0681 .0787
CEL .1123 .1011 .0962 .0929 .0891 .0610 .0892 .0825 .0792 .0852 .0816 .0953 .0719 .0747 .0747 .0783 .0817 .0783 .0781 .0999 .1112
ALP .1294 .1325 .1058 .0990 .0819 .0914 .1058 .0990 .1027 .1131 .1027 .1122 .0960 .0880 .0914 .0916 .0916 .0950 .0948 .0956 .1138
LUG .1297 .1364 .1060 .0993 .0955 .0776 .0986 .0919 .0886 .0919 .0886 .0985 .0854 .0810 .0776 .0846 .0846 .0812 .0810 .0915 .0956
BIT .1149 .1404 .0885 .0886 .0887 .0812 .0988 .0956 .0958 .1095 .1027 .1238 .0856 .0744 .0812 .0848 .0882 .0848 .0846 .1030 .1143
MEL .1221 .1325 .0885 .0921 .0819 .0812 .0919 .0852 .0850 .0952 .0919 .1122 .0784 .0711 .0812 .0814 .0882 .0848 .0846 .0886 .0958
BIV .1182 .1286 .0849 .0885 .0749 .0776 .0849 .0782 .0852 .0954 .0921 .1124 .0751 .0676 .0776 .0778 .0846 .0812 .0810 .0850 .0995
RIG .1291 .1322 .0952 .0988 .0886 .0878 .1021 .0954 .0990 .1023 .0990 .1266 .0888 .0776 .0878 .0880 .0880 .0914 .0912 .0919 .1030
KOU .1219 .1286 .0917 .0919 .0851 .0844 .0986 .0919 .0956 .0988 .0956 .1089 .0785 .0743 .0844 .0846 .0914 .0880 .0878 .0885 .0995
PLK .1369 .1326 .1024 .0922 .0955 .0881 .1059 .0991 .0959 .1026 .1029 .1306 .0926 .0813 .0915 .0917 .0952 .0952 .0950 .0991 .1104
VOL .1182 .1213 .0849 .0816 .0817 .0743 .0917 .0850 .0818 .0919 .0921 .1160 .0785 .0676 .0776 .0778 .0846 .0812 .0810 .0919 .1030
SPP .1257 .1288 .0919 .0886 .0887 .0812 .0988 .0921 .0888 .0990 .0993 .1233 .0856 .0744 .0846 .0848 .0916 .0882 .0880 .0956 .1067
LOE .1219 .1249 .0883 .0850 .0851 .0776 .0952 .0885 .0852 .0954 .0956 .1195 .0820 .0709 .0810 .0812 .0880 .0846 .0844 .0919 .1030
PUN .1219 .1249 .0883 .0850 .0851 .0776 .0952 .0885 .0852 .0954 .0956 .1195 .0820 .0709 .0810 .0812 .0880 .0846 .0844 .0919 .1030
OGO LOU HER AXD PRI ALH CAL AUS CEL ALP LUG BIT MEL BIV RIG KOU PLK VOL SPP LOE
LOU .0250
HER .0751 .0820
EXD .0443 .0508 .0820
PRI .0542 .0542 .0852 .0124
ALH .0815 .0883 .1171 .0849 .0746
CAL .0890 .0925 .1176 .0999 .0962 .0283
AUS .0883 .0883 .1100 .0747 .0645 .0218 .0479
CEL .0923 .0958 .1174 .0959 .0853 .0155 .0251 .0379
ALP .0847 .0776 .1281 .0776 .0741 .0711 .0821 .0677 .0715
LUG .0986 .0845 .1207 .0914 .0810 .0543 .0751 .0443 .0546 .0540
BIT .0988 .0984 .1317 .0848 .0846 .0577 .0821 .0511 .0648 .0476 .0412
MEL .0881 .0946 .1174 .0709 .0741 .0610 .0821 .0479 .0648 .0506 .0410 .0251
BIV .0811 .0909 .1135 .0674 .0706 .0609 .0820 .0478 .0646 .0473 .0409 .0282 .0062
RIG .0913 .0944 .1243 .0775 .0807 .0676 .0820 .0543 .0714 .0505 .0474 .0314 .0124 .0155
KOU .0948 .0978 .1171 .0741 .0773 .0642 .0854 .0510 .0680 .0473 .0377 .0282 .0093 .0124 .0155
PLK .0916 .0981 .1137 .0777 .0809 .0644 .0891 .0545 .0682 .0539 .0508 .0379 .0187 .0218 .0218 .0218
VOL .0845 .0909 .1135 .0674 .0706 .0576 .0785 .0445 .0613 .0473 .0409 .0282 .0093 .0062 .0155 .0124 .0155
SPP .0915 .0980 .1174 .0743 .0775 .0610 .0821 .0479 .0648 .0539 .0442 .0282 .0093 .0124 .0155 .0124 .0155 .0062
LOE .0879 .0944 .1135 .0708 .0740 .0576 .0785 .0445 .0613 .0505 .0409 .0250 .0062 .0093 .0124 .0093 .0124 .0031 .0031
PUN .0709 .0810 .0812 .0880 .0846 .0844 .0919 .1030 .0879 .0944 .1135 .0708 .0740 .0576 .0785 .0445 .0613 .0505 .0409 .0250
DEL, Fundulopanchax deltaense; WAL, Fp. walkeri; EXU, Aphyosemion exiguum; CYA, A. cyanostictum; LAB, A. labarrei; MIM, A. mimbon; MAC, A. maculatum; CAM, A. cameronense; OCC, A.
occelatum; AUR, A. aureum; CIT, A. citrineipinnis; COE, A. coeleste; LAM, A. lamberti; CHR, A. christyi; COG, A. cognatum; DEC, A. decorsei; ELE, A. elegans; MLP, A. melanopteron; STR, A. striatum;
GAB, A. gabunense; OGO, A. ogoense; LOU, A. louessense; HER, A. hera; EXD, A. exigoideum; PRI, A. primigenium; ALH, A. ahli; CAL, A. calliurum; AUS, A. australe; CEL, A. celiae; ALP, A.
(Chrom.) alpha; LUG, A. (Chrom.) lugens; BIT, A. (Chrom.) bitaeniatum; MEL, A. (Chrom.) melanogaster; BIV, A. (Chrom.) bivittatum; RIG, A. (Chrom.) riggenbachi; KOU, A. (Chrom.) kouamense;
PLK, A. (Chrom.) poliaki; VOL, A. (Chrom.) volcanum; SPP, A. (Chrom.) splendopleure; LOE, A. (Chrom.) loennbergii; PUN, A. (Chrom.) punctulatum. Abbreviations for species of the subgenus Chromaphyosemion
and distance between these species are in bold format
Fig. 2. Consensus tree based on distance method, maximum likelihood, maximum parsimony, and Bayesian inference methods. Numbers above the
branches are percentages of bootstrap values based on 500 replicates for each method (except for Bayesian inferences where the numbers represent the
percentage of trees with best likelihood scores in which the particular node was found). Values below 50% are not shown. Fundulopanchax species (a
monophyletic genus according to Murphy and Collier (1999)) were used to root the trees. Chromaphyosemion species formed a monophyletic group
(bootstrap values ranged from 67 to 75%). Aphyosemion (except A. hera) and Fundulopanchax sequences used are the same than those used in Murphy and
Collier (1999) and are available in GenBank. CAL, calliurum group; CAM, cameronense group; COL, coeleste group; ELE, elegans group; EXI, exiguum
group; GEO, georgiae group; STR, striatum group; UG, ungrouped species.
Fig. 3. Consensus tree based on distance method, maximum likelihood, maximum parsimony, and Bayesian inference methods. Numbers above or below
the branches are percentages of bootstrap values based on 500 replicates for each method (except for Bayesian inferences where the numbers represent the
percentage of trees with best likelihood scores in which the particular node was found). Only bootstrap values with maximum parsimony method are given
for intraspecies nodes. Values below 50% are not shown. Aphyosemion ahli was used to root the diVerent networks. Numbers correspond to sample location
on the map (Fig. 1). When more than one specimen were sequenced from the same location, they have been noted a, b, c, etc. References of specimens
collected at location 53 where two diVerent species have been identiWed are underlined.
Fig. 4. Scatter plot of the number of chromosomes versus the percentage
of bi-armed chromosomes for the 17 Chromaphyosemion karyotypes that
have been studied (Scheel, 1990; Völker et al., 2005). Species identiWcations
for Scheel (1974, 1990) karyotypes do not necessarily follow this
author as taxonomy has changed since 1990. In most of the cases, it was
possible to precisely determine what species Scheel (1974, 1990) did karyotype.
When it was not possible to unambiguously determine the species,
data are presented with a question mark. ALP, A. alpha; BIT, A. bitaeniatum;
BIV, A. bivittatum; KOU, A. kouamense; LUG, A. lugens; RIG, A.
riggenbachi; LOE, A. loennbergii; LOE?, doubtful status (A. splendopleure
?); MEL, A. melanogaster; PLK, A. poliaki; SPP, A. splendopleure; SP?,
undescribed species from Rio Muni (not necessarily identical to GEMHS
00/32); VOL, A. volcanum. The equation of the trend line is
yD¡0.091x + 1.9091. There is a statistically signiWcant relationship
(Spearman test) between these two variables.
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