364:, or terrestrial jelly nests. Most cophylines have very simple advertisement calls, consisting of single melodious notes that are repeated after regular intervals and for long periods of time, usually lasting several minutes. Correlated to the reproductive mode of the various cophyline lineages is their arboreal versus terrestrial or fossorial ecology, and apparently, multiple evolutionary shifts between arboreal and terrestrial habits have occurred in this subfamily.
53:
31:
388:, is not closely related to these two subfamilies, so microhylids colonised Madagascar at least twice. It is not however clear which subfamily is most closely related to Cophylinae+Scaphiophryninae, in part because the topology of deep nodes of the microhylid evolutionary tree is not satisfactorily resolved and remains unstable.
395:
in the rainforests of northern
Madagascar, and today only few species are known from more arid areas in the drier west of the island. As its sister subfamily Scaphiophryninae also has most of its centre of diversity in this area, it is probable that these subfamilies originated in northern Madagascar
404:
in ecologically-related morphological traits among the frogs, which led also to taxonomic confusion when only sparse morphological data were available; a problem that has been mostly but not fully rectified by the increased availability of DNA sequence data. Among the repeated
477:
Scherz, Mark D.; Vences, Miguel; Rakotoarison, Andolalao; Andreone, Franco; Köhler, Jörn; Glaw, Frank; Crottini, Angelica (2016). "Reconciling molecular phylogeny, morphological divergence and classification of
Madagascan narrow-mouthed frogs (Amphibia: Microhylidae)".
399:
The
Cophylinae are highly ecologically labile over evolutionary time: in several instances, phylogenetic relationships based on DNA sequence data clearly indicate repeated transitions from e.g. terrestrial to arboreal habits and back. This has resulted in extensive
583:
Rakotoarison, A.; Scherz, M.D.; Glaw, F.; Köhler, J; Andreone, F.; Franzen, M.; Glos, J.; Hawlitschek, O.; Jono, T.; Mori, A.; Ndriantsoa, S.H.; Raminosoa
Rasoamampionona, N.; Riemann, J.C.; Rödel, M.-O.; Rosa, G.M.; Vieites, D.R.; Crottini, A.; Vences, M. (2017).
1048:
Wollenberg, Katharina C.; Vieites, David R.; Van Der
Meijden, Arie; Glaw, Frank; Cannatella, David C.; Vences, Miguel (2008-08-01). "Patterns of Endemism and Species Richness in Malagasy Cophyline Frogs Support a Key Role of Mountainous Areas for Speciation".
204:
research has revealed a significant taxonomic gap in this subfamily, and an estimated 70+ candidate species were identified. Many of these have subsequently been described, as well as numerous new discoveries (e.g. 26 species of
359:
Cophylines are characterized by a derived mode of larval development: whereas most microhylids have a specialized filter-feeding tadpole, cophylines have non-feeding tadpoles that develop either in tree holes, terrestrial foam
396:
and radiated thence outward across the island. It remains to be seen if the description of the 70+ undescribed candidate species of
Cophylinae will influence this pattern once described and incorporated into spatial analyses.
1135:
661:
384:, another Madagascar-endemic subfamily. Thus, two subfamily units of Microhylidae are endemic to and probably originated on the island of Madagascar. What is also clear is that the third Madagascar-endemic subfamily,
1187:
Scherz, Mark D.; Hutter, Carl R.; Rakotoarison, Andolalao; Riemann, Jana C.; Rödel, Mark-Oliver; Ndriantsoa, Serge H.; Glos, Julian; Hyde
Roberts, Sam; Crottini, Angelica (2019-03-27). Crowther, Mathew S. (ed.).
848:
Peloso, Pedro L.V.; Frost, Darrel R.; Richards, Stephen J.; Rodrigues, Miguel T.; Donnellan, Stephen; Matsui, Masafumi; Raxworthy, Cristopher J.; Biju, S.d.; Lemmon, Emily
Moriarty (2016-04-01).
425:. The relationships of these frogs within the subfamily indicate that at least four different groups within this subfamily have independently evolved to be among the smallest frogs in the world.
641:
Blommers-Schlösser RMA (1975). "Observations on the larval development of some
Malagasy frogs, with notes on their ecology and biology (Anura: Dyscophinae, Scaphiophryninae and Cophylinae)".
1190:"Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera"
200:. Members of this subfamily range from minute (< 10 mm adult body size) to fairly large (> 100 mm adult body size), and they are highly ecologically diverse.
710:
de Sá, Rafael O.; Streicher, Jeffrey W.; Sekonyela, Relebohile; Forlani, Mauricio C.; Loader, Simon P.; Greenbaum, Eli; Richards, Stephen; Haddad, Célio F. B. (2012-12-10).
1369:
1412:
800:"Nuclear gene phylogeny of narrow-mouthed toads (Family: Microhylidae) and a discussion of competing hypotheses concerning their biogeographical origins"
1356:
1402:
910:"Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary"
908:
Feng, Yan-Jie; Blackburn, David C.; Liang, Dan; Hillis, David M.; Wake, David B.; Cannatella, David C.; Zhang, Peng (2017-07-03).
850:"The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae)"
586:"Describing the smaller majority: Integrative fast-track taxonomy reveals twenty-six new species of tiny microhylid frogs (genus
409:
that have evolved is miniaturised body size. In 2019, five new species of miniaturised frogs were described, including the genus
380:
have been a matter of some debate, and only recently has a tentative consensus emerged that they are most closely related to the
1407:
1304:
1361:
446:
1136:"Recurrent ecological adaptations revealed through a molecular analysis of the secretive cophyline frogs of Madagascar"
662:"Recurrent ecological adaptations revealed through a molecular analysis of the secretive cophyline frogs of Madagascar"
524:
Vieites, David R.; Wollenberg, Katharina C.; Andreone, Franco; Köhler, Jörn; Glaw, Frank; Vences, Miguel (2009-05-19).
712:"Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera"
52:
1374:
798:
van der
Meijden, Arie; Vences, Miguel; Hoegg, Simone; Boistel, Renaud; Channing, Alan; Meyer, Axel (2007-09-01).
1309:
978:
Brown, Jason L.; Sillero, Neftali; Glaw, Frank; Bora, Parfait; Vieites, David R.; Vences, Miguel (2016-01-06).
317:
39:
1257:
295:
232:
1330:
1201:
1150:
991:
921:
811:
723:
676:
487:
401:
1397:
526:"Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory"
129:
1082:
887:
47:
376:, as they are restricted to the island. Their affinities with other subfamilies of the diverse
1317:
1237:
1219:
1166:
1104:
record at Tsingy de Bemaraha confirms the presence of cophyline frogs in western Madagascar".
1074:
1066:
1027:
1009:
957:
939:
879:
871:
827:
759:
741:
692:
565:
547:
503:
1227:
1209:
1158:
1134:
Andreone, Franco; Vences, Miguel; Vieites, David R.; Glaw, Frank; Meyer, Axel (2005-02-01).
1113:
1058:
1017:
999:
947:
929:
861:
819:
749:
731:
684:
601:
555:
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392:
381:
1295:
312:
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274:
242:
1205:
1154:
995:
925:
815:
799:
727:
680:
491:
30:
1232:
1189:
1022:
979:
952:
909:
754:
711:
560:
525:
345:
331:
265:
152:
217:
As of December 2019, the following genera are recognised in the subfamily Cophylinae:
1391:
1086:
1062:
406:
299:
228:
201:
891:
415:
377:
326:
182:
114:
1322:
1214:
1004:
1343:
1289:
1162:
823:
688:
499:
421:
385:
289:
249:
1280:
1117:
373:
303:
236:
189:
138:
1223:
1070:
1013:
943:
875:
745:
736:
551:
934:
542:
340:
178:
94:
64:
1241:
1170:
1078:
1031:
961:
883:
831:
763:
696:
569:
507:
1274:
606:
585:
84:
980:"Spatial Biodiversity Patterns of Madagascar's Amphibians and Reptiles"
260:
222:
193:
146:
1348:
866:
849:
197:
74:
1251:
1100:
Andreone, Franco; Randrianirina, Jasmin E. (2008). "An unexpected
1335:
451:
Amphibian Species of the World: an Online Reference. Version 6.0
361:
185:
104:
1255:
621:
660:
Andreone F; Vences M; Vieites DR; Glaw F; Meyer M (2005).
372:
There is little doubt that the Cophylinae originated on
298:, Riemann, Rödel, Ndriantsoa, Glos, Roberts, Crottini,
1264:
782:Monograph of the frogs of the family Microhylidae
914:Proceedings of the National Academy of Sciences
530:Proceedings of the National Academy of Sciences
784:. Trustees of the British Museum, London, UK.
8:
1252:
472:
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29:
20:
1231:
1213:
1021:
1003:
951:
933:
865:
753:
735:
605:
559:
541:
440:
438:
434:
1182:
1180:
1143:Molecular Phylogenetics and Evolution
1129:
1127:
1043:
1041:
973:
971:
903:
901:
843:
841:
804:Molecular Phylogenetics and Evolution
793:
791:
669:Molecular Phylogenetics and Evolution
480:Molecular Phylogenetics and Evolution
7:
775:
773:
519:
517:
453:. American Museum of Natural History
14:
1413:Taxa named by Edward Drinker Cope
391:The subfamily Cophylinae has its
1063:10.1111/j.1558-5646.2008.00420.x
51:
1:
1215:10.1371/journal.pone.0213314
1005:10.1371/journal.pone.0144076
413:, and members of the genera
1403:Endemic fauna of Madagascar
1163:10.1016/j.ympev.2004.10.013
824:10.1016/j.ympev.2007.02.008
689:10.1016/j.ympev.2004.10.013
500:10.1016/j.ympev.2016.04.019
1429:
1118:10.11646/zootaxa.1812.1.2
445:Frost, Darrel R. (2016).
165:
160:
144:
137:
48:Scientific classification
46:
37:
28:
23:
737:10.1186/1471-2148-12-241
716:BMC Evolutionary Biology
935:10.1073/pnas.1704632114
543:10.1073/pnas.0810821106
447:"Cophylinae Cope, 1889"
40:Anodonthyla boulengerii
16:Subfamily of amphibians
780:Parker, H. W. (1934).
1408:Amphibian subfamilies
368:Origins and Evolution
607:10.3897/vz.67.e31595
239:& Crottini, 2016
235:, Andreone, Köhler,
209:described in 2017).
1206:2019PLoSO..1413314S
1155:2005MolPE..34..315A
996:2016PLoSO..1144076B
926:2017PNAS..114E5864F
920:(29): E5864–E5870.
816:2007MolPE..44.1017V
728:2012BMCEE..12..241D
681:2005MolPE..34..315A
492:2016MolPE.100..372S
393:centre of diversity
594:Vertebrate Zoology
590:) from Madagascar"
192:. It has over 100
1385:
1384:
1258:Taxon identifiers
867:10.1111/cla.12118
536:(20): 8267–8272.
349:
335:
321:
307:
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172:
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1420:
1378:
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1217:
1184:
1175:
1174:
1140:
1131:
1122:
1121:
1097:
1091:
1090:
1057:(8): 1890–1907.
1045:
1036:
1035:
1025:
1007:
975:
966:
965:
955:
937:
905:
896:
895:
869:
845:
836:
835:
810:(3): 1017–1030.
795:
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382:Scaphiophryninae
344:
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294:Scherz, Hutter,
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278:
264:
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226:
128:
56:
55:
33:
21:
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1303:
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1200:(3): e0213314.
1186:
1185:
1178:
1138:
1133:
1132:
1125:
1099:
1098:
1094:
1047:
1046:
1039:
990:(1): e0144076.
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370:
357:
313:Plethodontohyla
275:Madecassophryne
215:
156:
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127:
50:
17:
12:
11:
5:
1426:
1424:
1416:
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1327:
1314:
1301:
1286:
1270:
1268:
1262:
1261:
1256:
1248:
1247:
1176:
1149:(2): 315–322.
1123:
1092:
1037:
967:
897:
860:(2): 113–140.
837:
787:
769:
702:
675:(2): 315–322.
652:
633:
613:
600:(3): 271–398.
575:
513:
464:
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432:
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427:
369:
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356:
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13:
10:
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2:
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1229:
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1216:
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1203:
1199:
1195:
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1183:
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1177:
1172:
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1111:
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623:
622:"AmphibiaWeb"
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367:
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36:
32:
27:
22:
19:
1265:
1197:
1193:
1146:
1142:
1109:
1105:
1102:Rhombophryne
1101:
1095:
1054:
1050:
987:
983:
917:
913:
857:
853:
807:
803:
781:
719:
715:
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672:
668:
655:
646:
642:
636:
625:. Retrieved
616:
597:
593:
587:
578:
533:
529:
483:
479:
455:. Retrieved
450:
420:
416:Rhombophryne
414:
410:
398:
390:
378:Microhylidae
371:
358:
350:(41 species)
339:
327:Rhombophryne
325:
322:(11 species)
311:
296:Rakotoarison
288:
273:
270:(21 species)
259:
256:(12 species)
254:Müller, 1892
248:
233:Rakotoarison
221:
216:
206:
174:
173:
166:
145:
124:
115:Microhylidae
38:
18:
1344:iNaturalist
1290:Wikispecies
486:: 372–381.
422:Anodonthyla
386:Dyscophinae
336:(9 species)
308:(3 species)
279:Guibé, 1974
250:Anodonthyla
202:DNA barcode
188:endemic to
121:Subfamily:
24:Cophylinae
1398:Cophylinae
1392:Categories
1310:Cophylinae
1296:Cophylinae
1266:Cophylinae
854:Cladistics
722:(1): 241.
643:Beaufortia
627:2019-03-28
429:References
374:Madagascar
190:Madagascar
183:microhylid
175:Cophylinae
139:Type genus
125:Cophylinae
1224:1932-6203
1112:: 46–48.
1087:205781693
1071:1558-5646
1051:Evolution
1014:1932-6203
944:0027-8424
876:1096-0031
746:1471-2148
588:Stumpffia
552:0027-8424
407:ecomorphs
402:homoplasy
341:Stumpffia
318:Boulenger
283:monotypic
243:monotypic
207:Stumpffia
196:in eight
179:subfamily
167:See text
71:Kingdom:
65:Eukaryota
1281:Q2019009
1275:Wikidata
1242:30917162
1194:PLOS ONE
1171:15619444
1079:18485110
1032:26735688
984:PLOS ONE
962:28673970
892:84925667
884:34732021
832:17369057
764:23228209
697:15619444
570:19416818
508:27085671
346:Boettger
332:Boettger
266:Boettger
227:Scherz,
153:Boettger
111:Family:
95:Amphibia
85:Chordata
81:Phylum:
75:Animalia
61:Domain:
1336:2810016
1233:6436692
1202:Bibcode
1151:Bibcode
1106:Zootaxa
1023:4703303
992:Bibcode
953:5530686
922:Bibcode
812:Bibcode
755:3561245
724:Bibcode
677:Bibcode
649:: 7–26.
561:2688882
488:Bibcode
355:Biology
261:Cophyla
223:Anilany
194:species
161:Genera
147:Cophyla
101:Order:
91:Class:
1375:165822
1362:661434
1349:428527
1240:
1230:
1222:
1169:
1085:
1077:
1069:
1030:
1020:
1012:
960:
950:
942:
890:
882:
874:
830:
762:
752:
744:
695:
568:
558:
550:
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457:10 May
348:, 1881
334:, 1880
320:, 1882
306:, 2019
302:&
300:Vences
268:, 1880
229:Vences
213:Genera
198:genera
155:, 1880
132:, 1889
1139:(PDF)
1083:S2CID
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665:(PDF)
362:nests
186:frogs
177:is a
105:Anura
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459:2016
419:and
411:Mini
304:Glaw
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237:Glaw
130:Cope
1331:EoL
1323:JDB
1318:CoL
1305:ASW
1228:PMC
1210:doi
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1018:PMC
1000:doi
948:PMC
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