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Amblyrhynchus cristatus BELL, 1825

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Higher TaxaIguanidae, Iguania, Sauria, Squamata (lizards)
SubspeciesAmblyrhynchus cristatus cristatus BELL 1825
Amblyrhynchus cristatus godzilla MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017
Amblyrhynchus cristatus hassi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus hayampi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017
Amblyrhynchus cristatus jeffreysi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017
Amblyrhynchus cristatus mertensi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus nanus GARMAN 1892
Amblyrhynchus cristatus sielmanni EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus trillmichi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017
Amblyrhynchus cristatus venustissimus EIBL-EIBESFELDT 1956
Amblyrhynchus cristatus wikelskii MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017 
Common NamesE: Marine Iguana
G: Meerechse
albemarlensis: Isabela Marine Iguana
hassi: Santa Cruz Marine Iguana
mertensi: San Cristóbal Marine Iguana
nanus: Genovesa Marine Iguana
sielmanni: Pinta Marine Iguana
venustissimus: Española Marine Iguana 
SynonymAmblyrhynchus cristatus BELL 1825: 206
Iguana (A. [mblyrhynchus]) Cristatus — GRAY 1831: 37
Iguana (A. [mblyrhynchus]) Ater GRAY 1831
Amblyrhincus [sic] cristatus — DUMÉRIL & BIBRON 1837: 195
Amblyrhincus [sic] ater — DUMÉRIL & BIBRON 1837: 196
? Amblyrhincus [sic] Demarlii DUMÉRIL & BIBRON 1837: 197
Hypsilophus (Amblyrhynchus) cristatus — FITZINGER 1843: 55
Hypsilophus (Amblyrhynchus) ater — FITZINGER 1843: 55
Oreocephalus cristatus — GRAY 1845: 189
Amblyrhynchus cristatus — STEINDACHNER 1876
Amblyrhynchus cristatus — BOULENGER 1885: 185
Amblyrhynchus cristatus — SWASH & STILL 2000

Amblyrhynchus cristatus cristatus BELL 1825
Amblyrhynchus cristatus BELL 1825: 206
Amblyrhynchus cristatus albemarlensis EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus albemarlensis — BUCKLEY et al. 2016
Amblyrhynchus cristatus cristatus — BUCKLEY et al. 2016

Amblyrhynchus cristatus godzilla MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017

Amblyrhynchus cristatus hassi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus hassi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus hassi — BUCKLEY et al. 2016

Amblyrhynchus cristatus hayampi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017

Amblyrhynchus cristatus jeffreysi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017

Amblyrhynchus cristatus mertensi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus mertensi EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus mertensi — BUCKLEY et al. 2016

Amblyrhynchus cristatus nanus GARMAN 1892
Amblyrhynchus nanus GARMAN 1892
Amblyrhynchus cristatus nanus — BUCKLEY et al. 2016

Amblyrhynchus cristatus sielmanni EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus sielmanni EIBL-EIBESFELDT 1962
Amblyrhynchus cristatus sielmanni — BUCKLEY et al. 2016

Amblyrhynchus cristatus trillmichi MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017

Amblyrhynchus cristatus venustissimus EIBL-EIBESFELDT 1956
Amblyrhynchus cristatus venustissimus EIBL-EIBESFELDT 1956
Amblyrhynchus cristatus venustissimus — BUCKLEY et al. 2016

Amblyrhynchus cristatus wikelskii MIRALLES, MACLEOD, RODRÍGUEZ, IBÁÑEZ, JIMÉNEZ-UZCATEGUI, QUEZADA, VENCES & STEINFARTZ 2017 
DistributionGalapagos (Ecuador)

cristatus: Isla Fernandina (= Narborough I.)

albemarlensis: Isla Isabela (= Albemarle I.)

ater: Isla Pinzón (= Duncan I.)

hassi: Isla Santa Cruz (= Intefadigable I.); Type locality: ‘Indefatigable Südküste, westliche Akademiebucht (...) Indefatigable, Galápagos Inseln’. [ = South coast of Isla Santa Cruz, western part of Academy Bay].

hayampi: Galapagos Islands (Marchena and very likely its satellite islets); Type locality: NW coast of Isla Marchena.

godzilla: Galapagos Islands (Northern and north-eastern part of San Cristóbal and the satellite islets off the northern coast of San Cristóbal); Type locality: Galápagos archipelago, San Cristóbal island, Salinas colony (at Punta Pitt, SRS), at geographical coordinates (decimal format) 0.697247S, 89.27084W

jeffreysi: Galapagos Islands (Wolf, Darwin (tentative), Roca Redonda and very likely their satellite islets); Type locality: Isla Wolf, no exact locality.

mertensi: Isla San Cristóbal (= Chatham I.), Isla Santiago (= James I.); Type locality: ‘etwa 3 km südwest- lich der Wrack-Bucht der Insel Chatham, Galápagos Inseln’ [ = Isla San Cristóbal, about 3 km SW to Wreck Bay (vicinity of Puerto Baquerizo Moreno)].

nanus: Isla Genovesa (= Tower I.); Type locality: ‘Tower Island’ [ = Isla Genovesa].

sielmanni: Isla Pinta (= Abington I.); Type locality: ‘Westküste der Insel Abington’ [ = Isla Pinta, W coast].

trillmichi: Galapagos Islands (Santa Fé and very likely its satellite islets); Type locality: Galápagos archipelago, Santa Fé island, West coast, El Miedo, geographical coordinates (in decimal format) 0.82581S, 90.02861W.

venustissimus: Isla Española (= Hood I.), Gardener Island, Floreana, Champion; Type locality: ‘Nordküste der Insel Hood’ [ = Isla Española, N coast].

wikelskii: Galapagos Islands (Santiago, Rábida and very likely their satellite islets); Type locality: Crab-point, S of James Bay (Isla Santiago)

Type locality: restricted to (Eibl-Eibesfeldt 1956) Narborough (Fernandina).  
ReproductionHybridization: Hybrids between Amblyrhynchus cristatus and Conolophus subcristatus have been reported by several authors (Lücker & Feiler 2002). Amblyrhynchus cristatus, Conolophus subcristatus and Conolophus marthae can hybridize and produce viable offspring (Tzika et al. 2008). 
TypesHolotype: OUM (= OUMNH) 6176, Oxford University Museum of Natural History 6176
Neotype: SMF 64179 [Iguana ater Gray] (designated by Miralles et al. 2017), also the holotype of A. cristatus albemarlensis Eibl-Eibesfeldt, 1962; the latter nomen therefore becomes an objective junior synonym of Iguana ater Gray, 1831.
Holotype: BMNH 1946.8.30.20 (formerly numbered BMNH 99.5.4); syntype: FMNH 16946 [nanus]
Holotype: SMF 64179 [albemarlensis]
Holotype: SMF 57407 [hassi]
Holotype: SMF 57430 [mertensi]
Holotype: SMF 57417 [sielmanni]
Holotype: CAS 12471, coll. by J.R. Slevin and E.S. King in 1906 [hayampi]
Holotype: CAS 12690, coll. by J.R. Slevin in 1906 [jeffreysi]
Holotype: VCCDRS 2967, coll. by A. Macleod in March 2013 [godzilla]
Holotype: VCCDRS 3036, coll. by A. Ibáñez on 10 January 2016 [trillmichi]
Holotype: SMF 49851 [venustissimus]
Holotype: SMF 57447 (erroneously given as SFM), coll. by I. Eibl-Eibesfeldt in 1957, (specimen originally designated as paratype of A. c. mertensi); Paratypes: SMF (given as SFM) 57446, 57448, 57449, same data as holotype (all of them originally designated as paratypes of A. c. mertensi) [wikelskii] 
DiagnosisDiagnosis (genus): differs from all other iguanid genera in possessing the following combination of character states: a) posterolateral processes on the ventral surface of the premaxilla absent, b) anterior surface of premaxillary rostral body nearly flat, c) nasal process of the premaxilla nearly vertical, d) nasal capsule greatly inflated and nasal bones relatively large, e) frontal develops deep paired pockets on ventral surface, f) dorsal orbital borders wedge-shaped, g) maxilla flares outward below row of supralabial foramina, h) lacrimal very small, i) ventral process of squamosal large, j) anterodorsal surface of septomaxilla bears pronounced longitudinal crest, k) parasphenoid rostrum very short, l) stapes relatively thick, m) dorsal edge of dentary much higher than dorsal edge of surangular on either side of coronoid, n) anterior inferior alveolar foramen located between the coronoid and the splenial, not contacting the dentary, o) angular process of prearticular remains relatively small throughout ontogeny, p) crowns of posterior marginal teeth tricuspid, q) secondary cusps of tricuspid marginal teeth relatively large, only slightly smaller than the apical cusp, r) second ceratobranchials of hyoid apparatus separated from one another medially for most or all of their lengths, s) scapular fenestrae small or absent, t) posterior process of interclavicle does not extend beyond lateral corners of sternum, u) lateral processes of the interclavicle form angles of between 75° and 90° with the posterior process (interclavicle roughly T-shaped), v) sternal fontanelle small or absent, w) xiphisterna separated from one another medially, x) separable skull osteoderms develop over frontal, prefrontal, and nasal bones, y) superciliary scales quadrangular and non-overlapping, z) gular fold weakly developed, aa) digits of manus and pes partially webbed (John Iverson et al., pers. comm.). 
CommentSynonymy: Miralles et al. 2017 synonymized A. c. albemarlensis with A. c. cristatus.

Terra typica: Mexico (in error) [BELL 1825].

Habitat: partly marine.

Type species: Amblyrhynchus cristatus BELL 1825 is the type species of the genus Amblyrhynchus BELL 1825. 
EtymologyA. c. godzilla refers to the fictional saurian monster from the eponym movie franchise created by Tomoyuki Tanaka (IMDb, 2016). Explicitly mentioned in the title sequence of the 1990s American remake (Emmerich, 1998), the physical appearance and swimming behaviour of marine iguanas were a significant source of inspiration to the creature ́s designer (Patrick Tatopoulos, personal communication). The name is an invariable noun in apposition.

A. c. hayampi refers to the word used to designate the common iguana in Kichwa (a Quechuan language representing one of the three official languages of Ecuador and currently spoken by a million people in South America).

A. c. jeffreysi was named after Jeffreys Málaga, a ranger of the Galápagos National Park with an outstanding interest and enthusiasm for the fauna and flora of the Galápagos, and especially for marine iguanas. Jeffreys Málaga has been awarded several times ‘Guardaparque del Año,’ or Park Warden of the Year and has been an essential part of our team during several excursions to the Galápagos.

A. c. trillmichi was named after Prof. Dr. Fritz Trillmich, a foremost biological researcher in Germany who has spent a lifetime working on the biology of the Galápagos islands. For many years Fritz Trillmich worked together with Martin Wikelski on the island of Santa Fé, where their research on the local population produced important insights into the behaviour and ecology of marine iguanas. In addition to undertaking his own research, Fritz Trillmich is Professor Emeritus of Behavioural Ecology at the University of Bielefeld where he continues to inspire and mentor generations of young researchers who will continue his legacy for years to come.

A. c. wikelskii was named after Dr. Martin Wikelski in recognition of his outstanding research on marine iguanas for several decades. Martin Wikelski has been fascinated by marine iguanas, which served as a study system for his behavioural and ecological research from his PhD onwards. He has published important papers on marine iguana ecology and conservation, for instance demonstrating the devastating effects of an oil spill on this species, and showing that marine iguanas can shrink in response to starvation induced by El Niño–based climate fluctuations. 
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