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1

Berry, Kate A., Juan Pablo Muñoz-Pérez, Cristina P. Vintimilla-Palacios, and Christofer J. Clemente. "Morphological and performance modifications in the world’s only marine lizard, the Galápagos marine iguana, Amblyrhynchus cristatus." Biological Journal of the Linnean Society 133, no. 1 (March 20, 2021): 68–80. http://dx.doi.org/10.1093/biolinnean/blab002.

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Abstract The Galápagos marine iguana (Amblyrhynchus cristatus), the world’s only extant marine lizard, may have had one of the most unique and challenging transitions to aquatic life. Curiously, previous studies have identified relatively few physiological adaptations in the marine iguana; however, little is known about the extent of morphological specialization and performance trade-offs associated with the marine environment. By examining the morphology and locomotory performance of the marine iguana in comparison to their closely related mainland ancestors, the black spiny-tailed iguana (Ctenosaura similis) and green iguana (Iguana iguana), we found variation reflected specialization to ecological niches. However, variation was more pronounced among subspecies of marine iguana, suggesting that little morphological or performance modification is required for iguanids to successfully invade aquatic environments, thus raising the question why there are so few extant aquatic reptilian lineages. Our findings indicate that specialization for the marine environment likely resulted in a trade-off in sprint speed in a terrestrial environment, which may explain why other lizards have not undergone transitions to the marine environment. Additionally, we found that the magnitude of morphological and performance variation was more pronounced between subspecies of marine iguana than between iguanid species. This illustrates that the form-function relationship is more complex than previously thought and sheds light on the ecomorphological mysteries of the marine iguana.
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2

Ibáñez, Alejandro, Markus Menke, Galo Quezada, Gustavo Jiménez-Uzcátegui, Stefan Schulz, and Sebastian Steinfartz. "Diversity of compounds in femoral secretions of Galápagos iguanas (genera:AmblyrhynchusandConolophus), and their potential role in sexual communication in lek-mating marine iguanas (Amblyrhynchus cristatus)." PeerJ 5 (August 17, 2017): e3689. http://dx.doi.org/10.7717/peerj.3689.

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BackgroundChemical signals are widely used in the animal kingdom, enabling communication in various social contexts, including mate selection and the establishment of dominance. Femoral glands, which produce and release waxy secretions into the environment, are organs of central importance in lizard chemical communication. The Galápagos marine iguana (Amblyrhynchus cristatus) is a squamate reptile with a lek-mating system. Although the lekking behaviour of marine iguanas has been well-studied, their potential for sexual communication via chemical cues has not yet been investigated. Here we describe the diversity of the lipophilic fraction of males’ femoral gland secretions among 11 island populations of marine iguanas, and compare it with the composition of its sister species, the Galápagos land iguana (Conolophus subcristatus). We also conducted behavioural observations in marine iguana territorial males in order to explore the possible function of these substances in the context of male dominance in leks.MethodsFemoral secretions were analysed by gas chromatography coupled to mass spectrometry (GC–MS), and chromatography with a flame ionisation detector (GC-FID) in order to characterise the lipophilic composition. To understand the potential role of femoral secretions in marine iguana intraspecific communication, territorial males were sampled for their femoral glands and monitored to record their head bob rate—a territorial display behaviour in males—as well as the number of females present in their leks.ResultsWe found that the gland secretions were composed of ten saturated and unsaturated carboxylic acids ranging in chain length between C16and C24, as well as three sterols. Cholesterol was the main compound found. Intriguingly, land iguanas have a higher diversity of lipophilic compounds, with structural group of lipids (i.e. aldehydes) entirely absent in marine iguanas; overall the chemical signals of both species were strongly differentiated. Lipid profiles also differed among populations of marine iguanas from different islands, with some islands demonstrating a high diversity of lipophilic compounds (i.e. full spectra of compounds), and others lacking one or more compounds. Among the compounds most frequently found missing were 11- and 13-eicosenoic acids. Gland secretions of males with a better body condition and with a higher dominance status (i.e. those accompanied by females and with higher head bob display) were proportionately richer in C20-unsaturated fatty acids (11-eicosenoic acid).DiscussionLand and marine iguanas strongly diverged in their chemical composition of the femoral glands likely due to ecological differences between both species. Despite that marine iguana populations varied in their femoral gland composition that was not related to their genetic structure. Our results indicated that 11-eicosenoic acid may play an important role in intraspecific chemical communication in marine iguanas.
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3

FRICK, MICHAEL G. "Misconceptions about the ability of researchers to relocate the holotype of the Galapagos pink land iguana through the use of a passive integrated transponder." Zootaxa 2694, no. 1 (December 2, 2010): 57. http://dx.doi.org/10.11646/zootaxa.2694.1.4.

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Isla Isabela is the largest island in the Galapagos archipelago and hosts three endemic iguanid lizard species (Tzika et al. 2008, Gentile & Snell 2009): Amblyrhynchus cristatus (Bell, 1825), Conolophus subcristatus (Gray, 1831) and Conolophus marthae Gentile & Snell, 2009. The first iguanid listed exists as a marine species and the latter two are restricted to the terrestrial environment. It should be noted, however, that all three species can hybridize and produce viable offspring (Tzika et al. 2008). The International Union for Conservation of Nature (IUCN) lists the marine iguana A. cristatus as a “threatened” species and the land iguana species C. subcristatus as “vulnerable”. Currently there are no studies that assess the status of the iguana C. marthae, so its status is given as “not evaluated” by the IUCN.
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4

Scheibel, Jessica, Joan Garcia-Porta, Galo Quezada, and Alejandro Ibáñez. "Phylogeography and Prevalence of Hemoparasites (Apicomplexa: Eucoccidiorida) in Galápagos Marine Iguanas, Amblyrhynchus cristatus (Reptilia: Iguanidae)." Animals 12, no. 9 (April 28, 2022): 1142. http://dx.doi.org/10.3390/ani12091142.

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Parasitism is among the most common forms of coexistence of organisms of different species. Hemoparasites live in the bloodstream of the host where they complete different life-cycle stages. Members of the phylum Apicomplexa constitute a large portion of all hemoparasites infecting reptiles and their parasite transmitting vectors, including arthropods. In this study, we carried out a survey and molecular identification of hemoparasites in blood samples of the iconic Galápagos marine iguana (Amblyrhynchus cristatus). Major island populations of marine iguanas were sampled to examine large-scale biogeographic patterns of parasite diversity and prevalence. Nested PCRs were used to amplify segments of the 18S rRNA-gene of hemoparasites. Furthermore, ticks attached to marine iguanas were collected and analyzed in the same way to assess their potential use as a non-invasive method for the detection of hemoparasites in vertebrate host species. PCR products were sequenced and a phylogenetic analysis was carried out showing the presence of two genetically distinct clusters of hemoparasites, one more commonly distributed than the other one, belonging to the genera Hepatozoon and/or Hemolivia (Apicomplexa: Eucoccidiorida). Overall, 25% of marine iguanas were infected by hemoparasites. However, infection rates varied strongly among particular island populations (from 3.45% to 50%). Although marine iguanas are an extremely mobile species that has colonized all islands in the Galápagos archipelago, parasite occurrence was not related to geographical distance, suggesting that dispersal behavior has a minor role in parasite transmission. On most islands, females tended to have higher infection rates than males, but this relationship was only significant on one island. Overall, ticks and marine iguanas had similar prevalence and diversity of parasites. However, the infection profiles of ticks and their corresponding hosts (marine iguanas) did not mirror one another, indicating that this method cannot be used reliably to assess marine iguana infection status. Interestingly, we found that hemoparasite prevalence in marine iguanas and ticks tended to be positively correlated across islands. Our results indicate that certain populations of marine iguanas may have special mechanisms and adaptations to cope with parasite infection. In addition, other factors such as vector density, anthropogenic-related activities or the immunological state of marine iguanas could potentially affect the striking variation in hemoparasite prevalence across island populations.
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5

Olson, Storrs L. "The early scientific history of Galapagos iguanas." Archives of Natural History 41, no. 1 (April 2014): 141–53. http://dx.doi.org/10.3366/anh.2014.0217.

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The oldest known specimen of Galapagos marine iguana (Amblyrhynchus cristatus), now in the University Museum, Oxford, was originally thought to have come from Mexico. A plausible history of its origin with sealers in the Galapagos Islands about 1824 and transportation to and across Mexico is advanced. The naturalists David Douglas and John Scouler, on James (Santiago) Island in January 1825, encountered and attempted unsuccessfully to preserve specimens of the land iguana (Conolophus subcristatus) but only a Scouler specimen of marine iguana made it back to England, and it has since disappeared. Published and previously unpublished journal entries from the voyage of HMS Blonde, which had shore parties at Albemarle (Isabela) and Narborough (Fernandina) islands in March 1825, establish that the specimens on which the original description of Amblyrhynchus (later Conolophus) subcristatus J. E. Gray, 1831 , was based originated in the voyage of the Blonde. Banks Bay, Albemarle Island, is here designated as the type locality for Conolophus subcristatus. Specimens of the marine iguana were also brought back by the Blonde. The published accounts of Scouler and the voyage of the Blonde established the Galapagos as the true home of the marine iguana well before the return of Charles Darwin and HMS Beagle in 1836.
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6

Stekolnikov, Alexandr A. "Three new species of chigger mites (Acariformes: Trombiculidae) from the Galápagos Islands." Systematic and Applied Acarology 26, no. 2 (January 25, 2021): 325–42. http://dx.doi.org/10.11158/saa.26.2.1.

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Three new species of chigger mites are described from two species of iguana endemic to the Galápagos Islands. Odontacarus cruzi sp. nov. and Schoengastia galapa sp. nov. are found on the marine iguana, Amblyrhynchus cristatus Bell, and Eutrombicula pachytrichia sp. nov. is found on the Santa Fe land iguana, Conolophus pallidus Heller. All these mite species were collected, identified, and named by P.H. Vercammen-Grandjean, but remained undescribed until the present.
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7

Trillmich, Krisztina G. K., and Fritz Trillmich. "Foraging strategies of the marine iguana, Amblyrhynchus cristatus." Behavioral Ecology and Sociobiology 18, no. 4 (February 1986): 259–66. http://dx.doi.org/10.1007/bf00300002.

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8

MacLeod, Amy, Ariel Rodríguez, Miguel Vences, Pablo Orozco-terWengel, Carolina García, Fritz Trillmich, Gabriele Gentile, Adalgisa Caccone, Galo Quezada, and Sebastian Steinfartz. "Hybridization masks speciation in the evolutionary history of the Galápagos marine iguana." Proceedings of the Royal Society B: Biological Sciences 282, no. 1809 (June 22, 2015): 20150425. http://dx.doi.org/10.1098/rspb.2015.0425.

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The effects of the direct interaction between hybridization and speciation—two major contrasting evolutionary processes—are poorly understood. We present here the evolutionary history of the Galápagos marine iguana ( Amblyrhynchus cristatus ) and reveal a case of incipient within-island speciation, which is paralleled by between-island hybridization. In-depth genome-wide analyses suggest that Amblyrhynchus diverged from its sister group, the Galápagos land iguanas, around 4.5 million years ago (Ma), but divergence among extant populations is exceedingly young (less than 50 000 years). Despite Amblyrhynchus appearing as a single long-branch species phylogenetically, we find strong population structure between islands, and one case of incipient speciation of sister lineages within the same island—ostensibly initiated by volcanic events. Hybridization between both lineages is exceedingly rare, yet frequent hybridization with migrants from nearby islands is evident. The contemporary snapshot provided by highly variable markers indicates that speciation events may have occurred throughout the evolutionary history of marine iguanas, though these events are not visible in the deeper phylogenetic trees. We hypothesize that the observed interplay of speciation and hybridization might be a mechanism by which local adaptations, generated by incipient speciation, can be absorbed into a common gene pool, thereby enhancing the evolutionary potential of the species as a whole.
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9

Trillmich, Krisztina G. K. "The Mating System of the Marine Iguana (Amblyrhynchus cristatus)1." Zeitschrift für Tierpsychologie 63, no. 2-3 (April 26, 2010): 141–72. http://dx.doi.org/10.1111/j.1439-0310.1983.tb00084.x.

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10

Colosimo, Giuliano, Gabriele Gentile, Carlos A. Vera, Christian Sevilla, Glenn P. Gerber, Hans D. Westermeyer, and Gregory A. Lewbart. "Conservation of a flagship species: Health assessment of the pink land iguana, Conolophus marthae." PLOS ONE 17, no. 3 (March 29, 2022): e0257179. http://dx.doi.org/10.1371/journal.pone.0257179.

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The pink land iguana, Conolophus marthae, is one of four species of iguanas (three terrestrial and one marine) in the Galápagos Islands, and the only one listed as critically endangered by the IUCN. The species can only be found on the north-west slopes of the highest volcano on Isabela Island and was first described to science in 2009. As part of a population telemetry study, a health assessment was authorized by the Galápagos National Park. Wild adult iguanas were captured on Wolf Volcano in September 2019 and April 2021 to record morphological and physiological parameters including body temperature, heart rate, intraocular pressures, tear formation, and infrared iris images. Blood samples were also collected and analyzed. An i-STAT portable blood analyzer was used to obtain values for base excess in the extracellular fluid compartment (BEecf), glucose (Glu), hematocrit (HctPCV), hemoglobin (Hb), ionized calcium (iCa), partial pressure of carbon dioxide (pCO2), partial pressure of oxygen (pO2), percent oxygen saturation (sO2%), pH, potassium (K), and sodium (Na). When possible, data were compared to previously published and available data for the other Galápagos iguanas. The results reported here provide baseline values that will be useful in detecting changes in health status among pink land iguanas affected by climate change, invasive species, anthropogenic threats, or natural disturbances. The collected data also provide an invaluable resource for conservation scientists planning to implement conservation strategies, like translocations, that may temporarily alter these baseline values.
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11

Rauch, Norbert. "Competition of Marine Iguana Females (Amblyrhynchus Cristatus) for Egg-Laying Sites." Behaviour 107, no. 1-2 (1988): 91–105. http://dx.doi.org/10.1163/156853988x00214.

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12

Tompkins, Paul, and Matthias Wolff. "Galápagos macroalgae: A review of the state of ecological knowledge." Revista de Biología Tropical 65, no. 1 (September 23, 2016): 375. http://dx.doi.org/10.15517/rbt.v65i1.18139.

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Previous work has highlighted the critical role of macroalgal productivity and dynamics in supporting and structuring marine food webs. Spatio-temporal variability in macroalgae can alter coastal ecosystems, a relationship particularly visible along upwelling-influenced coastlines. As a result of its equatorial location and nutrient rich, upwelling-influenced waters, the Galápagos Archipelago in the East Pacific, hosts a productive and biodiverse marine ecosystem. Reports and collections of macroalgae date back to the Beagle voyage, and since then, more than three hundred species have been reported. However, their ecology and functional role in the ecosystem is not well understood. According to various disparate and in part anecdotal sources of information, abundant and diverse communities exist in the Western regions of the archipelago, the North is essentially barren, and in the central/South abundance and distribution is variable and less well defined. Both oceanographic conditions and herbivore influence have been theorized to cause this pattern. Extensive changes in macroalgal productivity and community composition have occurred during strong ENSO events, and subsequent declines in marine iguana (an endemic and iconic grazer) populations have been linked to these changes. Iguanas are only one species of a diverse and abundant group of marine grazers in the system, highlighting the potentially important role of macroalgal productivity in the marine food web. This review represents a first compilation and discussion of the available literature and presents topics for future research.
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MacLeod, Amy, Volker Koch, Carolina García-Parra, Fritz Trillmich, and Sebastian Steinfartz. "New highly polymorphic microsatellite loci for the Galápagos marine iguana, Amblyrhynchus cristatus." Amphibia-Reptilia 33, no. 3-4 (2012): 533–36. http://dx.doi.org/10.1163/15685381-00002854.

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We describe the development and characterisation of six new dinucleotide motif microsatellite loci for populations of marine iguanas (Amblyrhynchus cristatus), endemic to the Galápagos archipelago. Primers were based on microsatellite-bearing sequences and initially developed using universally labelled primers. When analysed across 5 populations (representing 150 individuals), new loci displayed, on average, high levels of genetic diversity (range: 2-13 alleles, mean: 5.73) and values of heterozygosity (range: 0.0-0.906, mean: 0.605). No consistent deviations from Hardy-Weinberg equilibrium or significant linkage disequilibrium were observed, and all loci were shown to be free of common microsatellite errors. Utilising the 13 previously available microsatellite loci for this species, we describe here four multiplex combinations for the successful amplification of 19 microsatellite loci for marine iguanas. This powerful set of highly polymorphic markers will allow researchers to explore future questions regarding the ecology, evolution, and conservation of this unique species.
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14

Christman, J. E., K. J. Lohmann, M. Hirschfeld, J. P. Muñoz-Pérez, J. Garcia, J. A. Hernandez, H. Westermeyer, and G. A. Lewbart. "Intraocular pressure using rebound tonometry in the Galápagos marine iguana (Amblyrhynchus cristatus)." International Journal of Veterinary Research 3, no. 1 (January 1, 2023): 1–11. http://dx.doi.org/10.52547/injvr.3.1.1.

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Wikelski, Martin, Victor Carrillo, and Fritz Trillmich. "ENERGY LIMITS TO BODY SIZE IN A GRAZING REPTILE, THE GALAPAGOS MARINE IGUANA." Ecology 78, no. 7 (October 1997): 2204–17. http://dx.doi.org/10.1890/0012-9658(1997)078[2204:eltbsi]2.0.co;2.

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16

Vitousek, Maren N., James S. Adelman, Nathan C. Gregory, and James J. H. St Clair. "Heterospecific alarm call recognition in a non-vocal reptile." Biology Letters 3, no. 6 (October 2, 2007): 632–34. http://dx.doi.org/10.1098/rsbl.2007.0443.

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The ability to recognize and respond to the alarm calls of heterospecifics has previously been described only in species with vocal communication. Here we provide evidence that a non-vocal reptile, the Galápagos marine iguana ( Amblyrhynchus cristatus ), can eavesdrop on the alarm call of the Galápagos mockingbird ( Nesomimus parvulus ) and respond with anti-predator behaviour. Eavesdropping on complex heterospecific communications demonstrates a remarkable degree of auditory discrimination in a non-vocal species.
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Rassmann, K., F. Trillmich, and D. Tautz. "Hybridization between the Galápagos land and marine iguana (Canolophus subcristatusandAmblyrhynchus cristatus) on Plaza Sur." Journal of Zoology 242, no. 4 (August 1997): 729–39. http://dx.doi.org/10.1111/j.1469-7998.1997.tb05822.x.

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18

STEINFARTZ, SEBASTIAN, and ADALGISA CACCONE. "A set of highly discriminating microsatellite loci for the Galapagos marine iguana Amblyrhynchus cristatus." Molecular Ecology Notes 6, no. 3 (September 2006): 927–29. http://dx.doi.org/10.1111/j.1471-8286.2006.01402.x.

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19

French, Susannah S., Dale F. DeNardo, Timothy J. Greives, Christine R. Strand, and Gregory E. Demas. "Human disturbance alters endocrine and immune responses in the Galapagos marine iguana (Amblyrhynchus cristatus)." Hormones and Behavior 58, no. 5 (November 2010): 792–99. http://dx.doi.org/10.1016/j.yhbeh.2010.08.001.

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20

Tellkamp, Frederik, Franziska Lang, Alejandro Ibáñez, Lena Abraham, Galo Quezada, Stefan Günther, Mario Looso, et al. "Proteomics of Galápagos Marine Iguanas Links Function of Femoral Gland Proteins to the Immune System." Molecular & Cellular Proteomics 19, no. 9 (June 24, 2020): 1523–32. http://dx.doi.org/10.1074/mcp.ra120.001947.

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Communication between individuals via molecules, termed chemosignaling, is widespread among animal and plant species. However, we lack knowledge on the specific functions of the substances involved for most systems. The femoral gland is an organ that secretes a waxy substance involved in chemical communication in lizards. Although the lipids and volatile substances secreted by the femoral glands have been investigated in several biochemical studies, the protein composition and functions of secretions remain completely unknown. Applying a proteomic approach, we provide the first attempt to comprehensively characterize the protein composition of femoral gland secretions from the Galápagos marine iguana. Using samples from several organs, the marine iguana proteome was assembled by next-generation sequencing and MS, resulting in 7513 proteins. Of these, 4305 proteins were present in the femoral gland, including keratins, small serum proteins, and fatty acid-binding proteins. Surprisingly, no proteins with discernible roles in partner recognition or inter-species communication could be identified. However, we did find several proteins with direct associations to the innate immune system, including lysozyme C, antileukoproteinase (ALP), pulmonary surfactant protein (SFTPD), and galectin (LGALS1) suggesting that the femoral glands function as an important barrier to infection. Furthermore, we report several novel anti-microbial peptides from the femoral glands that show similar action against Escherichia coli and Bacillus subtilis such as oncocin, a peptide known for its effectiveness against Gram-negative pathogens. This proteomics data set is a valuable resource for future functional protein analysis and demonstrates that femoral gland secretions also perform functions of the innate immune system.
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Drent, J., W. D. Van Marken Lichtenbelt, and M. Wikelski. "Effects of foraging mode and season on the energetics of the Marine Iguana,Amblyrhynchus cristatus." Functional Ecology 13, no. 4 (August 1999): 493–99. http://dx.doi.org/10.1046/j.1365-2435.1999.00337.x.

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Rassmann, K., D. Tautz, F. Trillmich, and C. Gliddon. "The microevolution of the Galápagos marine iguana Amblyrhynchus cristatus assessed by nuclear and mitochondrial genetic analyses." Molecular Ecology 6, no. 5 (May 1997): 437–52. http://dx.doi.org/10.1046/j.1365-294x.1997.00209.x.

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Berger, Silke, Martin Wikelski, L. Michael Romero, Elisabeth K. V. Kalko, and Thomas Rödl. "Behavioral and physiological adjustments to new predators in an endemic island species, the Galápagos marine iguana." Hormones and Behavior 52, no. 5 (December 2007): 653–63. http://dx.doi.org/10.1016/j.yhbeh.2007.08.004.

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Thaller, Maria C., Marco Ciambotta, Manuela Sapochetti, Luciana Migliore, Whashington Tapia, Virna Cedeño, and Gabriele Gentile. "Uneven frequency of Vibrio alginolyticus-group isolates among different populations of Galápagos marine iguana (Amblyrhynchus cristatus)." Environmental Microbiology Reports 2, no. 1 (January 20, 2010): 179–84. http://dx.doi.org/10.1111/j.1758-2229.2009.00132.x.

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Wikelski, M., B. Gall, and F. Trillmich. "Ontogenetic changes in food intake and digestion rate of the herbivorous marine iguana (Amblyrhynchus cristatus, Bell)." Oecologia 94, no. 3 (June 1993): 373–79. http://dx.doi.org/10.1007/bf00317112.

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Chiari, Ylenia, Scott Glaberman, Pedro Tarroso, Adalgisa Caccone, and Julien Claude. "Ecological and evolutionary influences on body size and shape in the Galápagos marine iguana (Amblyrhynchus cristatus)." Oecologia 181, no. 3 (April 4, 2016): 885–94. http://dx.doi.org/10.1007/s00442-016-3618-1.

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Wikelski, Martin, and Fritz Trillmich. "Foraging Strategies of the Galapagos Marine Iguana (Amblyrhynchus Cristatus) : Adapting Behavioral Rules To Ontogenetic Size Change." Behaviour 128, no. 3-4 (1994): 255–79. http://dx.doi.org/10.1163/156853994x00280.

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Arrivillaga, Cristina, and Tom W. Brown. "Tail bifurcation in a Marine Iguana, Amblyrhynchus cristatus (Reptilia: Squamata: Iguanidae), from Isla Santa Cruz, Galápagos Islands." Reptiles & Amphibians 27, no. 3 (December 7, 2020): 415–18. http://dx.doi.org/10.17161/randa.v27i3.14858.

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Glaberman, Scott, Louis Du Pasquier, and Adalgisa Caccone. "Characterization of a Nonclassical Class I MHC Gene in a Reptile, the Galápagos Marine Iguana (Amblyrhynchus cristatus)." PLoS ONE 3, no. 8 (August 6, 2008): e2859. http://dx.doi.org/10.1371/journal.pone.0002859.

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30

Rauch, N. "Female habitat choice as a determinant of the reproductive success of the territorial male marine iguana (Amblyrhynchus cristatus)." Behavioral Ecology and Sociobiology 16, no. 2 (January 1985): 125–34. http://dx.doi.org/10.1007/bf00295146.

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Christensen, Kiana, and Keegan M. Melstrom. "Quantitative analyses of squamate dentition demonstrate novel morphological patterns." PLOS ONE 16, no. 9 (September 10, 2021): e0257427. http://dx.doi.org/10.1371/journal.pone.0257427.

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Squamates are ideal subjects for investigating relationships between diet and dental patterns because they exhibit wide dietary diversity, marked variation in dental shape, and are taxonomically abundant. Despite this, well-established links between diet and dental morphology are primarily qualitative in nature, with specific patterns of squamate dental complexity remaining largely unknown. Here, we use quantitative methods and a broad taxonomic dataset to quantify key patterns in squamate dental morphology, including re-examining the relationship between dentition and diet, testing for differences in complexity between dentigerous elements, and exploring the effect of ontogenetic dietary shifts in dental complexity in two iguanid genera. Our findings support previous research by demonstrating that species consuming more plant material possess more complex teeth. We did not find significant complexity differences between the left and right dentigerous elements nor the upper and lower jaws, with the exception of Amblyrhynchus cristatus, the marine iguana, which possesses significantly more complex dentary teeth than premaxillary and maxillary teeth. We find discordant patterns when testing for dental complexity changes through ontogeny. Amblyrhynchus, which is primarily herbivorous throughout its lifetime, increases dental complexity through ontogeny, whereas Ctenosaura, which is generally insectivorous as juveniles and herbivorous as adults, decreases dental complexity. Although preliminary, this research documents and quantifies novel patterns of squamate dental complexity and exhibits the possibilities for further research on the diversity of squamate dental morphology.
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MacLeod, Amy, and Sebastian Steinfartz. "The conservation status of the Galápagos marine iguanas, Amblyrhynchus cristatus: a molecular perspective." Amphibia-Reptilia 37, no. 1 (2016): 91–109. http://dx.doi.org/10.1163/15685381-00003035.

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Traditionally, conservation management focuses efforts on taxonomic units. However, when the taxa used do not reflect biologically meaningful units, such methods should be reconsidered to avoid the loss of irreplaceable biodiversity. The Galápagos marine iguana (Amblyrhynchus cristatus) is listed as Vulnerable on the IUCN Red List of Threatened Species and is facing growing anthropogenic threats. Currently, management is based on a taxonomy which is questionable in the light of recent molecular data. As such, there is a danger that evolutionarily significant populations may be left vulnerable to extirpation. Herein, we apply molecular data to elucidate the population structure of this species across the Galápagos archipelago, and thus advise conservation management in the absence of a revised taxonomy. Applying a wealth of molecular data including 12 microsatellite loci and 1181 bp of the mitochondrial control region in over 1200 individuals, we delineate distinct populations and prioritize their management. Bayesian population structure analysis revealed 10 distinct population clusters, which we propose as management units (MUs). All MUs are significantly differentiated, with one unit on San Cristóbal Island being particularly distinct in terms of both microsatellite loci and mitochondrial data. Based on estimates of the genetic effective population size (Ne), we find the MUs comprised of populations occurring on Floreana, Española, Marchena, and San Cristóbal to be alarmingly small. In consideration of both Neand anthropogenic threats, we recommend that conservation practitioners focus efforts on Floreana and San Cristóbal islands, and argue that better census size estimates of populations are urgently needed.
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Fields, Peter A., Chad M. Strothers, and Mark A. Mitchell. "Function of muscle-type lactate dehydrogenase and citrate synthase of the Galápagos marine iguana, Amblyrhynchus cristatus, in relation to temperature." Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology 150, no. 1 (May 2008): 62–73. http://dx.doi.org/10.1016/j.cbpb.2008.01.008.

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Hugi, Jasmina, and Marcelo R. Sánchez-Villagra. "Life History and Skeletal Adaptations in the Galapagos Marine Iguana (Amblyrhynchus cristatus) as Reconstructed with Bone Histological Data—A Comparative Study of Iguanines." Journal of Herpetology 46, no. 3 (September 2012): 312–24. http://dx.doi.org/10.1670/11-071.

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35

Klein, Nicole, Alexandra Houssaye, James M. Neenan, and Torsten M. Scheyer. "Long bone histology and microanatomy of Placodontia (Diapsida: Sauropterygia)." Contributions to Zoology 84, no. 1 (February 23, 2015): 59—S15. http://dx.doi.org/10.1163/18759866-08401005.

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Placodontia, an enigmatic group of durophagous and in part heavily armoured animals, were members of Sauropterygia, the most diverse and successful group of Mesozoic marine reptiles. Microanatomy and histology of long bones of several armoured and non-armoured Placodontia were studied, covering most of their taxonomic breadth, to elucidate the paleoecology, physiology, and lifestyle of its members. Results reveal an unexpected and not phylogenetically or stratigraphically related disparity of microanatomical and histological features for the group. The non-armoured Paraplacodus and the heavily armoured Psephoderma grew with lamellar-zonal bone tissue type, which is typical for modern sauropsids. In the former, the tissue is nearly avascular surrounding a compacted medullary region, whereas in the latter, the lamellar-zonal bone tissue is vascularized framing a large open medullary cavity and a perimedullary region. Armoured Henodus and Placodontia indet. aff. Cyamodus as well as non-armoured Placodus exhibit a reduced medullary cavity and grew with highly vascularized plexiform to radiating fibro-lamellar bone. Several long bones of Placodontia indet. show circumferential fibro-lamellar bone and can be distinguished into two groups on the basis of microanatomical features. In addition, all bones that grew with fibro-lamellar bone show locally primary spongeous-like architecture and had secondarily widened primary osteons throughout the cortex, resulting in a secondarily spongeous tissue. The highly vascularized fibro-lamellar bone of these Placodontia indicates growth rates comparable to that of open marine ichthyosaurs. Differences in microanatomy and bone histology as expressed by a principal component analysis, thus clearly indicate different paleoecologies, including differences in lifestyle and swimming modes and capabilities in Placodontia. This would have reduced competition in the shallow marine environments of the Tethys and might be a key to their success and diversity. A certain developmental plasticity among the studied placodonts is interpreted as response to different environmental conditions as is obvious from inter- and intraspecific histological variation. Most striking is the difference in life history strategy in armoured Psephoderma and non-armoured Paraplacodus when compared to armoured Henodus, Placodontia indet. aff. Cyamodus, non-armoured Placodus, and Placodontia indet. Bone tissue of Psephoderma and Paraplacodus indicates low growth rates and a low basal metabolic rate, as many modern sauropsids have such as the marine iguana, whereas the others grew with extremely fast growth rates, more typical for birds and mammals, indicating an increased basal metabolic rate.
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Luis A. Ramos-Vázquez, Nahíra Arocho-Hernández, Cielo Figuerola-Hernández, José L. Herrera-Giraldo, and Jan P. Zegarra-Vila. "Status of seabirds, habitat, and invasive species in the Cordillera Reef Nature Reserve, Puerto Rico." Journal of Caribbean Ornithology 34 (March 2, 2021): 1–11. http://dx.doi.org/10.55431/jco.2021.34.1-11.

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Abstract Seabird populations have been decreasing worldwide, and many are threatened or species of special concern. The Cordillera Reef Nature Reserve (CRNR) is located off the northeastern coast of Puerto Rico and is part of the Culebra Important Bird Area. We conducted a seabird population survey in April 2018 consisting of terrestrial point counts and boat surveys on Icacos, Ratones, Lobos, La Blanquilla, and Diablo cays, as well as one unnamed cay. The purpose of the survey was to document and count the various seabird species, describe the available nesting habitat, and document the presence of invasive vertebrates on the different cays of the CRNR. Of the 25 bird species documented throughout the CRNR, five were seabird species. Among these, Red-billed Tropicbirds (Phaethon aethereus) and Brown Boobies (Sula leucogaster) were the only species nesting within the CRNR, while Magnificent Frigatebirds (Fregata magnificens) and Brown Pelicans (Pelecanus occidentalis) flew over and roosted on the cays. We also detected a pair of Audubon’s Shearwaters (Puffinus lherminieri) in Ratones Cay, the first record of this species for the CRNR. Of all the assessed cays, Diablo provided the most suitable nesting habitat for Red-billed Tropicbirds and Brown Boobies. Among the cays, we documented invasive vertebrate species, such as black rats (Rattus rattus) and green iguanas (Iguana iguana) on Icacos and Ratones, but none on Diablo. This study highlights the significance of the CRNR as an important seabird area, adds new sightings to the available seabird information, documents the occurrence of invasive vertebrates, and reiterates the importance of monitoring seabird colonies for their conservation. Keywords Audubon's Shearwater, Brown Booby, Caribbean, habitat assessment, invasive species, Phaethon aethereus, Puerto Rico, Puffinus lherminieri, Red-billed Tropicbird, seabirds, Sula leucogaster Resumen Estado de las aves marinas, el hábitat y las especies invasoras en la Reserva Natural Arrecifes de la Cordillera, Puerto Rico • Las poblaciones de aves marinas han disminuido en todo el mundo y muchas especies están amenazadas o son de especial preocupación. La Reserva Natural Arrecifes de la Cordillera (RNAC) se encuentra en la costa nordeste de Puerto Rico y es parte de Culebra, un Área de Importancia para la Conservación de las Aves. En abril de 2018 se realizó un censo poblacional de aves marinas utilizando puntos de conteo en tierra y censos en bote en los cayos Icacos, Ratones, Lobos, La Blanquilla, Diablo y en un cayo sin nombre. El propósito de este censo fue documentar y cuantificar las especies de aves marinas presentes, describir el hábitat disponible para la nidificación y documentar la presencia de vertebrados invasores en los diferentes cayos de la RNAC. Se documentaron un total de 25 especies de aves en toda la reserva, de las cuales cinco fueron aves marinas. Phaethon aethereus y Sula leucogaster fueron las únicas especies que se encontraron nidificando, mientras que Fregata magnificens y Pelecanus occidentalis fueron observadas volando y descansando. Se detectó también la presencia de un par de invidivuos de Puffinus lherminieri en cayo Ratones, siendo el primer registro de esta especie para la RNAC. De todos los cayos evaluados, Cayo Diablo presentó el hábitat de nidificación más adecuado para Phaethon aethereus y Sula leucogaster. Se registraron, además, especies de vertebrados invasores como ratas negras (Rattus rattus) e iguanas verdes (Iguana iguana) en cayos Icacos y Ratones, pero ninguna en Cayo Diablo. Este estudio destaca la importancia de la RNAC como un área importante para la aves marinas; incluye nuevos avistamientos en la información disponible del grupo, documenta la presencia de vertebrados invasores y reitera la importancia del monitoreo de colonias de aves marinas para su conservación. Palabras clave aves marinas, Caribe, evaluación de hábitat, Phaethon aethereus, Puerto Rico, Puffinus lherminieri, Sula leucogaster, vertebrados invasores Résumé Statut des oiseaux marins, des habitats et des espèces envahissantes dans la Reserva Natural Arrecifes de la Cordillera, à Porto Rico • Les populations d’oiseaux marins ont décliné mondialement et beaucoup d’entre elles sont classées comme menacées ou préoccupantes. La Cordillera Reef Nature Reserve (CRNR) est située sur la côte nord-est de Porto Rico et est une Aire Importante pour les Oiseaux et la Biodiversité (Important Bird and Biodiversity Area). Un suivi de population d’oiseaux marins a été effectué en avril 2018, constistant à la réalisation de points d’écoute terrestres et de suivis maritimes par bateau sur les cayes d’Icacos, de Ratones, de Lobos, de La Blanquilla, de Diablo, et une caye sans nom. L’objectif de cette étude est d’y documenter and compter les différentes espèces d’oiseaux marins, de décrire l’habitat de nidification disponible et de documenter la présence d’espèces de vertébrés invasives dans les différentes cayes de la CRNR. Sur les 25 espèces d'oiseaux recensées dans tout le CRNR, cinq étaient des oiseaux marins. Seuls le Phaéton à bec rouge (Phaethon aethereus) et le Fou brun (Sula leucogaster) y ont niché alors que le Frégate superbe (Fregata magnificens) et le Pélican brun (Pelecanus occidentalis) ont été observés en vol et en dortoir dans la réserve. Nous avons aussi détecté la présence d’un couple de Puffin d’Audubon (Puffinus lherminieri) sur la caye de Ratones, la première observation de cette espèce pour la CRNR. De toutes les cayes évaluées, Diablo a montré l’habitat de nidification le plus adéquat pour le Phaéton à bec rouge et le Fou brun. La présence d’espèces invasives, comme le Rat noir (Rattus rattus) et l’Iguane vert (Iguana iguana) a été observée sur Icacos et Ratones, mais aucune n’a été détectée sur Diablo. Cette étude souligne l’importance de la CRNR comme une aire importante pour les oiseaux marins, met à jour les informations disponibles pour ces espèces marines, documente la présence de vertébrés invasifs et rappelle l’importance des suivis de colonies d’oiseaux marins pour leur conservation. Mots clés Caraïbes, espèces invasives, évaluation de l'habitat, fou brun, oiseaux marins, Phaethon aethereus, Phaéton à bec rouge, Porto Rico, Puffin d’Audubon, Puffinus lherminieri, Sula leucogaster
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37

Wikelski, M., L. M. Romero, and H. L. Snell. "Marine Iguanas Oiled in the Galapagos." Science 292, no. 5516 (April 20, 2001): 437c—438. http://dx.doi.org/10.1126/science.292.5516.437c.

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Wikelski, Martin, Vanessa Wong, Brett Chevalier, Niels Rattenborg, and Howard L. Snell. "Marine iguanas die from trace oil pollution." Nature 417, no. 6889 (June 2002): 607–8. http://dx.doi.org/10.1038/417607a.

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Wikelski, Martin, and Corinna Thom. "Marine iguanas shrink to survive El Niño." Nature 403, no. 6765 (January 2000): 37–38. http://dx.doi.org/10.1038/47396.

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40

Wikelski, Martin. "Evolution of body size in Galapagos marine iguanas." Proceedings of the Royal Society B: Biological Sciences 272, no. 1576 (August 24, 2005): 1985–93. http://dx.doi.org/10.1098/rspb.2005.3205.

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Body size is one of the most important traits of organisms and allows predictions of an individual's morphology, physiology, behaviour and life history. However, explaining the evolution of complex traits such as body size is difficult because a plethora of other traits influence body size. Here I review what we know about the evolution of body size in a group of island reptiles and try to generalize about the mechanisms that shape body size. Galapagos marine iguanas occupy all 13 larger islands in this Pacific archipelago and have maximum island body weights between 900 and 12 000 g. The distribution of body sizes does not match mitochondrial clades, indicating that body size evolves independently of genetic relatedness. Marine iguanas lack intra- and inter-specific food competition and predators are not size-specific, discounting these factors as selective agents influencing body size. Instead I hypothesize that body size reflects the trade-offs between sexual and natural selection. We found that sexual selection continuously favours larger body sizes. Large males establish display territories and some gain over-proportional reproductive success in the iguanas' mating aggregations. Females select males based on size and activity and are thus responsible for the observed mating skew. However, large individuals are strongly selected against during El Niño-related famines when dietary algae disappear from the intertidal foraging areas. We showed that differences in algae sward (‘pasture’) heights and thermal constraints on large size are causally responsible for differences in maximum body size among populations. I hypothesize that body size in many animal species reflects a trade-off between foraging constraints and sexual selection and suggest that future research could focus on physiological and genetic mechanisms determining body size in wild animals. Furthermore, evolutionary stable body size distributions within populations should be analysed to better understand selection pressures on individual body size.
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Rödl, Thomas, Silke Berger, L. Michael Romero, and Martin Wikelski. "Tameness and stress physiology in a predator-naive island species confronted with novel predation threat." Proceedings of the Royal Society B: Biological Sciences 274, no. 1609 (November 28, 2006): 577–82. http://dx.doi.org/10.1098/rspb.2006.3755.

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Tame behaviour, i.e. low wariness, in terrestrial island species is often attributed to low predation pressure. However, we know little about its physiological control and its flexibility in the face of predator introductions. Marine iguanas ( Amblyrhynchus cristatus ) on the Galápagos Islands are a good model to study the physiological correlates of low wariness. They have lived virtually without predation for 5–15 Myr until some populations were first confronted with feral cats and dogs some 150 years ago. We tested whether and to what extent marine iguanas can adjust their behaviour and endocrine stress response to novel predation threats. Here, we show that a corticosterone stress response to experimental chasing is absent in naive animals, but is quickly restored with experience. Initially, low wariness also increases with experience, but remains an order of magnitude too low to allow successful escape from introduced predators. Our data suggest that the ability of marine iguanas to cope with predator introductions is limited by narrow reaction norms for behavioural wariness rather than by constraints in the underlying physiological stress system. In general, we predict that island endemics show flexible physiological stress responses but are restricted by narrow behavioural plasticity.
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Wikelski, M. "Body Size, Performance and Fitness in Galapagos Marine Iguanas." Integrative and Comparative Biology 43, no. 3 (July 1, 2003): 376–86. http://dx.doi.org/10.1093/icb/43.3.376.

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43

Rubenstein, Dustin R., and Martin Wikelski. "Steroid hormones and aggression in female Galápagos marine iguanas." Hormones and Behavior 48, no. 3 (September 2005): 329–41. http://dx.doi.org/10.1016/j.yhbeh.2005.04.006.

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Wikelski, Martin, and Michaela Hau. "Is There an Endogenous Tidal Foraging Rhythm in Marine Iguanas?" Journal of Biological Rhythms 10, no. 4 (December 1995): 335–50. http://dx.doi.org/10.1177/074873049501000407.

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Wikelski, M., and Peter H. Wrege. "Niche expansion, body size, and survival in Galápagos marine iguanas." Oecologia 124, no. 1 (July 21, 2000): 107–15. http://dx.doi.org/10.1007/s004420050030.

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WIKELSKI, MARTIN, CHRIS CARBONE, and FRITZ TRILLMICH. "Lekking in marine iguanas: female grouping and male reproductive strategies." Animal Behaviour 52, no. 3 (September 1996): 581–96. http://dx.doi.org/10.1006/anbe.1996.0199.

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Richardson, Kristen M., John B. Iverson, and Carolyn M. Kurle. "Marine subsidies likely cause gigantism of iguanas in the Bahamas." Oecologia 189, no. 4 (March 8, 2019): 1005–15. http://dx.doi.org/10.1007/s00442-019-04366-4.

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Cooper, J. E., and W. A. Laurie. "Investigation of deaths in marine iguanas (Amblyrhynchus cristatus) on Galapagos." Journal of Comparative Pathology 97, no. 2 (March 1987): 129–36. http://dx.doi.org/10.1016/0021-9975(87)90032-6.

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Laurie, W. A., and D. Brown. "Population Biology of Marine Iguanas (Amblyrhynchus cristatus). III. Factors Affecting Survival." Journal of Animal Ecology 59, no. 2 (June 1990): 545. http://dx.doi.org/10.2307/4880.

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Dellinger, Thomas, and Gisela von Hegel. "Sex Identification through Cloacal Probing in Juvenile Marine Iguanas (Amblyrhynchus cristatus)." Journal of Herpetology 24, no. 4 (December 1990): 424. http://dx.doi.org/10.2307/1565065.

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