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Leiopelma; Temporal range: Miocene–Recent PreꞒ Ꞓ O S D C P T J K Pg N
	Hochstetter's frog (Leiopelma hochstetteri)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Amphibia
Order:	Anura
Suborder:	Archaeobatrachia
Family:	Leiopelmatidae; Mivart, 1869
Genus:	Leiopelma; Fitzinger, 1861
Species
	See text
	Distribution of Leiopelmatidae (in black)

Leiopelma is a genus of New Zealand primitive frogs, belonging to the suborder Archaeobatrachia. It is the only genus in the monotypic family Leiopelmatidae. The leiopelmatids' relatively basal form indicates they have an ancient lineage.^[1] While some taxonomists have suggested combining the North American frogs of the genus Ascaphus in the family Ascaphidae with the New Zealand frogs of the genus Leiopelma in the family Leiopelmatidae, the current consensus is that these two groups constitute two separate families.^[2]^[3] The three extant species of Leiopelmatidae are only found in New Zealand.^[4]

Overview

The New Zealand primitive frogs' defining characteristics are their extra vertebrae (for a total of nine) and the remains of the tail muscles (the tail itself is absent in adults, although it is present in the younger frogs, which need the extra skin surface until their lungs are fully developed). The family Ascaphidae (found only in North America), of the same suborder, shares these primitive characteristics, hence the two have often been described as related, or even part of the same family.

Late jump recovery is unique in Leiopelmatidae. When leiopelmatid species jump, they land in a "belly flop" fashion, repositioning their limbs for takeoff for the next jump only after hitting the ground with the ventral surface of their torsos. The appearance of early jump recovery in more advanced taxa is a key innovation in anuran evolution.^[5]

They are unusually small frogs, only 5 cm (2.0 in) in length. Most species lay their eggs in moist ground, typically under rocks or vegetation. After hatching, the tadpoles nest in the male's back, all without the need for standing or flowing water. However, Hochstetter's frog lays its eggs in shallow ponds and has free-living tadpoles, although they do not swim far from the place of hatching, or even feed, before metamorphosing into adult frogs.^[6] Lifespans may be long (more than 30 years) for such small organisms.^[7]

Introduced fauna are thought to have had a negative impact on these native frogs, with 93% of all reported predation events on native frogs being attributed to introduced fauna,^[8] primarily ship rats.

Taxonomy & systematics

Species

Family Leiopelmatidae

Genus Leiopelma
- Archey's frog, Leiopelma archeyi Turbott, 1942
- Hamilton's frog, Leiopelma hamiltoni McCulloch, 1919
- Hochstetter's frog, Leiopelma hochstetteri Fitzinger, 1861

Extinct species

Three extinct species are known by subfossil remains, also from New Zealand. They became extinct during the past 1,000 years.^[9]

†Aurora frog, Leiopelma auroraensis
†Markham's frog, Leiopelma markhami^[10]
†Waitomo frog, Leiopelma waitomoensis^[11]

One species from the late Pliocene period has recently been described.^[12]

†Leiopelma bishopi Easton, Tennyson and Rawlence, 2021

Two species are known from Miocene deposits of the Saint Bathans fauna, with indeterminate remains possibly representing additional species^[13]^[14]

†Leiopelma miocaenale
†Leiopelma acricarina

Evolutionary history

DNA analysis indicates that Leiopelmatidae share a distant common ancestry with Ascaphidae to the exclusion of all other frogs, and Leiopelmatidae and Ascaphidae diverged from all other frogs around 200 million years ago.^[15] L. archeyi and L. hochstetteri are thought to have diverged from each other between 40 and 50 million years ago, based on genomic divergence estimates. Fossils of the genus are known from the early Miocene (19–16 million years ago) aged St Bathans Fauna of New Zealand.^[13]

Anura

	Leiopelmatidae

	Ascaphidae

Bombianura

Costata

Bombinatoridae

	Alytidae

	Discoglossidae

Pipanura

Xenoanura

	Pipidae

	Rhinophrynidae

	Pelobatidae

	Megophryidae

	Sooglossidae

	Nasikabatrachidae

Notogaeanura

Australobatrachia

Calyptocephalellidae

	Myobatrachidae

	Limnodynastidae

Nobleoanura

Ranoides

References

^
- Roelants, Kim; Franky Bossuyt (February 2005). "Archaeobatrachian paraphyly and Pangaean diversification of crown-group frogs". Systematic Biology. 54 (1): 111–126. doi:10.1080/10635150590905894. PMID 15805014.
- San Mauro, Diego; Miguel Vences; Marina Alcobendas; Rafael Zardoya; Axel Meyer (May 2005). "Initial diversification of living amphibians predated the breakup of Pangaea" (PDF). American Naturalist. 165 (5): 590–599. doi:10.1086/429523. JSTOR 10.1086/429523. PMID 15795855. S2CID 17021360.
^ Frost, Darrel R. (2015). "Leiopelmatidae Mivart, 1869". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 1 September 2015.
^
- J.M. Conlon et al. / Peptides 30 (2009) 1069–1073
- Cannatella, David (2008). "Leiopelmatidae. Leiopelma". The Tree of Life Web Project. Retrieved 1 September 2015.
^ "DOC: Photo-stage and Archey's Frog". Retrieved 2005-12-05.
^ Essner, RL Jr; Suffian, DJ; Bishop, PJ; Reilly, SM (2010). "Landing in basal frogs: evidence of saltational patterns in the evolution of anuran locomotion". Naturwissenschaften. 97 (10): 935–9. Bibcode:2010NW.....97..935E. doi:10.1007/s00114-010-0697-4. PMID 20625697. S2CID 18602582.
^ Zweifel, Richard G. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. p. 85. ISBN 978-0-12-178560-4.
^ Bell, Ben D.; et al. (2004). "The fate of a population of the endemic frog Leiopelma pakeka (Anura: Leiopelmatidae) translocated to restored habitat on Maud Island, New Zealand". New Zealand Journal of Zoology. 31 (2): 123–131. doi:10.1080/03014223.2004.9518366.
^ Egeter, Bastian; Robertson, Bruce C.; Bishop, Phillip J. (2015). "A Synthesis of Direct Evidence of Predation on Amphibians in New Zealand, in the Context of Global Invasion Biology". Herpetological Review. 46: 512–519.
^
*Worthy, Trevor H. (1987). "Osteology of Leiopelma (Amphibia: Leiopelmatidae) and descriptions of three new subfossil Leiopelma species". Journal of the Royal Society of New Zealand. 17 (3): 201–251. doi:10.1080/03036758.1987.10418160.
- Worthy, Trevor H. (1987). "Palaeoecological information concerning members of the frog genus Leiopelma: Leiopelmatidae in New Zealand". Journal of the Royal Society of New Zealand. 17 (4): 409–420. doi:10.1080/03036758.1987.10426482.
- Nadia Webster (2004). "Native frog captive husbandry manual" (PDF). Department of Conservation. Archived from the original (PDF) on 4 March 2016. Retrieved 29 January 2015.
^ "Holotype of Leiopelma markhami". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 17 July 2010.
^ "Holotype of Leiopelma waitomoensis". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 17 July 2010.
^ Easton, Luke J.; Tennyson, Alan J. D.; Rawlence, Nicolas J. (2021-10-12). "A new species of Leiopelma frog (Amphibia: Anura: Leiopelmatidae) from the late Pliocene of New Zealand". New Zealand Journal of Zoology. 49 (3): 215–224. doi:10.1080/03014223.2021.1979053. ISSN 0301-4223. S2CID 243120256.
^ ^a ^b Worthy, Th; Tennyson, Ajd; Scofield, Rp; Hand, Sj (December 2013). "Early Miocene fossil frogs (Anura: Leiopelmatidae) from New Zealand". Journal of the Royal Society of New Zealand. 43 (4): 211–230. doi:10.1080/03036758.2013.825300. hdl:2328/35958. ISSN 0303-6758. S2CID 84562226.
^ Updating The Record from the Early Miocene St Bathans Fauna, Central Otago and its Significance for Documenting the Assembly of New Zealand’s Terrestrial Biota, Conference Paper · July 2014
^ Feng, Yan-Jie; Blackburn, David C.; Liang, Dan; Hillis, David M.; Wake, David B.; Cannatella, David C.; Zhang, Peng (2017-07-18). "Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary". Proceedings of the National Academy of Sciences. 114 (29): E5864 – E5870. doi:10.1073/pnas.1704632114. ISSN 0027-8424. PMC 5530686. PMID 28673970.