Project 291: T. J. Gaudin, R. J. Emry, J. R. Wible. 2009. The phylogeny of living and extinct pangolins (Mammalia, Pholidota) and associated taxa: a morphology based analysis. Journal of Mammalian Evolution. 16 (4):235-305.
Abstract
The present study was undertaken in order to effect a comprehensive phylogenetic analysis of the Order Pholidota, examining seven of the eight currently recognized extant species (absent is Manis culionensis, formerly recognized as a subspecies of Manis javanica) and nearly all the well-known fossil taxa, and employing a wide range of osteological characters from the entire skeleton. In addition, the relationship of pangolins to several putative early Tertiary relatives, including palaeanodonts and the enigmatic “edentate” Eurotamandua joresi, were investigated. The goal of the study was to improve understanding of the systematics and the biogeographic and evolutionary history of the pangolins. A computer-based cladistic analysis of phylogenetic relationships among seven extant species of pangolins, five extinct pangolin species (including all but one of the well-preserved taxa), as well as Eurotamandua and two genera of metacheiromyid palaeanodonts, Palaeanodon and Metacheiromys, was performed based upon 395 osteological characteristics of the skull and postcranial skeleton. Characters were polarized via comparison to the following successive outgroups: the basal feliform carnivoran Nandinia binotata and the hedgehog Erinaceus sp., a eulipotyphlan laursiatherian placental. A revised classification is presented based on the results of the analysis. The results support the monophyly of Pholidota and Palaeanodonta by providing new anatomical characters that can serve to diagnose a pangolin/palaeanodont clade, termed here Pholidotamorpha. Pholidota is defined so as to include all living and fossil pangolins, including all three taxa of middle Eocene “edentates” from the Messel fauna of Germany, among them Eurotamandua joresi. The results do not support the monophyly of the remaining two Messel “edentates” originally placed in the same genus Eomanis, which is restricted to the type species Eomanis waldi. Euromanis, new genus, is named with Eomanis krebsi Storch and Martin, 1994, as the type species, to form a new combination Euromanis krebsi (Storch and Martin, 1994). The analysis strongly supports the monophyly of a crown clade of pangolins diagnosed by many anatomical synapomorphies, the Superfamily Manoidea. This crown clade can in turn be divided into two monophyletic families: the Patriomanidae, which includes two Tertiary taxa, Patriomanis americana and Cryptomanis gobiensis, and the Manidae, which includes all the extant pangolins. The relationship of the Tertiary European pangolin Necromanis to these two families is unresolved. Within Manidae, the extant species are divided into three well-supported, monophyletic genera, Manis for the Asian pangolins, Smutsia for the African ground pangolins, and Phataginus for the African tree pangolins. The latter two form a monophyletic African assemblage, the subfamily Smutsiinae. The biogeographic implications of this phylogeny are examined. A European origin for Pholidota is strongly indicated. The fossil record of pangolins would seem to support a European origin for the modern forms, with subsequent dispersal into sub-Saharan African and then to southern Asia, and the phylogeny produced in this analysis is consistent with such scenario.Read the article »
Project DOI: 10.7934/P291, http://dx.doi.org/10.7934/P291
This project contains | Matrices |
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Download Project SDD File | Total scored cells: 5602 Total media associated with cells: 0 Total labels associated with cell media: 0 |
Characters | |
Total characters: 395 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 1013 Total character states with associated media: 113 Total character states with media with labels:113 Total unordered/ordered characters:395/0 |
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MorphoBank Project 291
MorphoBank Project 291
- Creation Date:
15 June 2009 - Publication Date:
15 June 2009 - Project views: 98402
- Matrix downloads: 40
This research
supported by
Authors' Institutions
- University of Tennessee
- Carnegie Museum of Natural History
- Smithsonian Institution
Members
member name | taxa | specimens | media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||||
Timothy Gaudin Project Administrator | 21 | 18 | 63 | 395 | 116 | 121 | 5602 (5602, 0, 0) | 0 | 0 | 0 | 0 | |||||
MorphoBank Curator Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 | |||||
Robert Emry Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 | |||||
John Wible Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M731' (17 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] Erinaceus sp. Last Modified in 12/05/13 | 28 | 369 | 367 | 0 | 0 | 0 | 0 | 4 |
[2] Nandinia binotata Last Modified in 12/05/13 | 11 | 386 | 384 | 0 | 0 | 0 | 0 | 4 |
[3] Manis crassicaudata Last Modified in 12/05/13 | 13 | 389 | 382 | 0 | 0 | 0 | 0 | 4 |
[4] Manis javanica Last Modified in 12/05/13 | 1 | 436 | 394 | 0 | 0 | 0 | 0 | 4 |
[5] Manis pentadactyla Last Modified in 12/05/13 | 6 | 417 | 389 | 0 | 0 | 0 | 0 | 4 |
[6] Smutsia gigantea Last Modified in 12/05/13 | 5 | 450 | 390 | 0 | 0 | 0 | 0 | 4 |
[7] Smutsia temminckii Last Modified in 12/05/13 | 8 | 405 | 387 | 0 | 0 | 0 | 0 | 4 |
[8] Phataginus tetradactyla Last Modified in 12/05/13 | 1 | 417 | 394 | 0 | 0 | 0 | 0 | 4 |
[9] Phataginus tricuspis Last Modified in 12/05/13 | 0 | 445 | 395 | 0 | 0 | 0 | 0 | 4 |
[10] Patriomanis americana Last Modified in 12/05/13 | 35 | 369 | 360 | 0 | 0 | 0 | 0 | 4 |
[11] Cryptomanis gobiensis Last Modified in 12/05/13 | 172 | 225 | 223 | 0 | 0 | 0 | 0 | 4 |
[12] Necromanis sp. Last Modified in 12/05/13 | 195 | 205 | 200 | 0 | 0 | 0 | 0 | 4 |
[13] Eomanis waldi Last Modified in 12/05/13 | 153 | 248 | 242 | 0 | 0 | 0 | 0 | 4 |
[14] Euromanis krebsi Last Modified in 12/05/13 | 309 | 89 | 86 | 0 | 0 | 0 | 0 | 4 |
[15] Eurotamandua joresi Last Modified in 12/05/13 | 179 | 221 | 216 | 0 | 0 | 0 | 0 | 4 |
[16] Palaeanodon sp. Last Modified in 12/05/13 | 186 | 214 | 209 | 0 | 0 | 0 | 0 | 4 |
[17] Metacheiromys sp. Last Modified in 12/05/13 | 79 | 317 | 316 | 0 | 0 | 0 | 0 | 4 |
Project views
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Total project views | 98402 | |
Project overview | 5766 | |
Matrix views | 2647 | Matrix landing page (2283 views); The phylogeny of living and extinct pangolins (Mammalia, Pholidota) and associated taxa: a morphology based analysis. (364 views); |
Media views | 56052 | Media search (5834 views); M32471 (829 views); M32480 (795 views); M32478 (814 views); M32470 (792 views); M32475 (984 views); M32473 (950 views); M32474 (792 views); M32481 (1022 views); M32489 (832 views); M32479 (823 views); M32529 (701 views); M32491 (808 views); M32497 (792 views); M32500 (781 views); M32493 (805 views); M32518 (703 views); M32503 (849 views); M32513 (792 views); M32482 (827 views); M32507 (870 views); M32508 (967 views); M32528 (694 views); M32502 (829 views); M32525 (778 views); M32483 (823 views); M32494 (857 views); M32472 (824 views); M32512 (782 views); M32531 (766 views); M32510 (781 views); M32520 (696 views); M32501 (815 views); M32511 (743 views); M32522 (675 views); M32504 (807 views); M32506 (906 views); M32519 (755 views); M32499 (785 views); M32514 (729 views); M32487 (849 views); M32527 (741 views); M32476 (835 views); M32521 (697 views); M32505 (792 views); M32477 (812 views); M32492 (855 views); M32532 (717 views); M32488 (831 views); M32530 (720 views); M32485 (792 views); M32524 (680 views); M32515 (816 views); M32495 (781 views); M32516 (755 views); M32496 (807 views); M32490 (825 views); M32509 (804 views); M32498 (834 views); M32523 (701 views); M32486 (826 views); M32517 (699 views); M32484 (783 views); M32526 (693 views); |
Specimen list | 19974 | |
Taxon list | 10860 | |
Bibliography | 1429 | |
Documents list | 1674 |
Project downloads
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 336 | |
Matrix downloads | 40 | The phylogeny of living and extinct pangolins (Mammalia, Pholidota) and associated taxa: a morphology based analysis. (40 downloads); |
Document downloads | 2 | Copyright information for exemplar media (2 downloads); |
Project downloads | 294 |