Comparative immunology of Galapagos iguana hemoglobins.

The antigenic properties of the major hemoglobin component of the Galapgaos iguanas were studied using second-approximation qualitative and quantitative immunochemical techniques. Phylogenetic distances, relative to the Galapagos marine iguana. Amblyrhynchus cristatus, were established on the basis of immunological cross-reactions.     

Variation among populations of Galapagos land iguanas (Conolophus): contrasts of phylogeny and ecology

A phylogenetic scheme derived via multivariate analyses of adaptively neutral scale characteristics is compared to patterns of ecological adaptation in body size and shape, hatchling size, clutch size, and reproductive seasonality, in extant populations of Galapagos land iguanas (genus Conolophus). Three groups of land iguana populations are identified, the oldest being the population of Isla Santa Fe, followed by the populations of the central islands (Santa Cruz, Plaza Sur and Baltra), the youngest populations are those of the western islands (Fernandina and Isabela). Patterns of ecological similarity among these populations are not concordant with phylogenetic lineage. Populations most similar in ecological characteristics are often phylogenetically divergent. Adaptation to local conditions by iguana populations is apparently more important than phylogenetic constraint in explaining variation in ecological characteristics. The assumption that phylogenetically closely-related organisms are also ecologically more similar than less closely-related organisms is not supported by this evidence. Some previous studies may have been misled by using ecological characteristics to derive phylogenetic lineages, resulting in circular support of the assumption.     

Biochemical and microbiological evidence for fermentative digestion in free-living land iguanas (Conolophus pallidus) and marine iguanas (Amblyrhynchus cristatus) on the Galápagos archipelago

Herbivorous lizards are potentially capable of high digestive efficiency, but the presence of an indigenous microbial population has been implied from measurements of activity rather than directly studied. This study is the first to provide direct biochemical and microbiological evidence for fermentative digestion in free-living land iguanas (Conolophus pallidus) and marine iguanas (Amblyrhynchus cristatus) from the Galapagos archipelago. In marine iguanas, the stomach and large capacious colon contained ca. 32% and 60%, respectively, of the weight of total gut content. Total volatile fatty acid concentration was ca. 150 and 180 mM, respectively, for marine and land iguanas. Molar proportions of acetate, propionate, and butyrate (80.3%, 9.5%, and 3.5%) in land iguana fecal samples were similar to those for marine iguanas. Examination of fecal samples using confocal and transmission electron microscopy, as well as cultivable counts, revealed a dense and diverse population of bacteria, with spores prominent. Total culturable counts of anaerobes (2.22x10(8) g(-1) wet weight of fecal material) outnumbered aerobes on average by a factor of ca. 700. Combined, these results strongly support the contention that these unique herbivorous lizards are largely dependent on the presence and metabolic activities of a resident bacterial population in order to hydrolyze and ferment plant polymers that are indigestible to the host.     

Development of primers to characterize the mitochondrial control region of Galápagos land and marine iguanas (Conolophus and Amblyrhynchus)

Primers were developed for the amplification and sequencing of the mitochondrial control region of Galápagos land (Conolophus) and marine (Amblyrhynchus) iguanas. Sequences were obtained for four land iguana samples from two islands and for 28 marine iguana samples from three islands. A series of 70–80 bp tandem repeats adjacent to the control region are described and preliminary quantification of intra‐ and interspecific sequence divergence is included.     

Species-specific evolution of class I MHC genes in iguanas (Order: Squamata; Subfamily: Iguaninae)

Over the last few decades, the major histocompatibility complex (MHC) has emerged as a model for understanding the influence of natural selection on genetic diversity in populations as well as for investigating the genetic basis of host resistance to pathogens. However, many vertebrate taxa remain underrepresented in the field of MHC research, preventing its application to studies of disease, evolution, and conservation genetics in these groups. This is particularly true for squamates, which are by far the most diversified order of non-avian reptiles but have not been the subject of any recent MHC studies. In this paper, we present MHC class I complementary DNA data from three squamate species in the subfamily Iguaninae (iguanas): the Galápagos marine iguana (Amblyrhynchus cristatus), the Galápagos land iguana (Conolophus subcristatus), and the green iguana (Iguana iguana). All sequences obtained are related to the few published class I genes from other squamates. There is evidence for multiple loci in each species, and the conserved alpha-3 domain appears to be evolving in a species-specific manner. Conversely, there is some indication of shared polymorphism between species in the peptide-binding alpha-1 and alpha-2 domains, suggesting that these two regions have different phylogenetic histories. The great similarity between alpha-3 sequences in marine iguanas in particular suggests that concerted evolution is acting to homogenize class I loci within species. However, while less likely, the data are also compatible with a birth and death model of evolution.     

Hybridization between the Galápagos land and marine iguana (Canolophus subcristatus and Amblyrhynchus cristatus) on Plaza Sur

Hybridization plays an important role in the evolution of some of the vertebrate taxa on the Galápagos Islands, such as the Darwin finches. Conversely, only a single possible hybrid between the Galápagos marine iguana ( Amblyrhynchus ) and the land iguana ( Conolophus ) has been reported from the island Plaza Sur. In this paper, the hybrid status of a morphologically unusual iguana from this island is confirmed, using restriction fragment length polymorphism (RFLP) analyses of the nuclear ribosomal DNA (rDNA). Sequencing of the hybrid's mitochondrial cytochrome b gene revealed that it was the offspring of a female land iguana and a male marine iguana. Preliminary molecular analyses of morphologically typical marine and land iguanas from Plaza Sur did not detect introgression of nuclear or mitochondrial markers between species. The potential significance of hybridization for the evolution of the Plaza Sur iguana populations is discussed.     

Influences of parasites and thermoregulation on grouping tendencies in marine iguanas

I determined whether grouping behavior influences parasite loadand body temperature of Galápagos marine iguanas, reptilesthat rest gregariously. Mobile (or predatory) Ornithodoros ticks(4.7 mm average body length) approached at a ground speed of65 cm/min and parasitized sleeping marine iguanas for 3.7 hper night, drawing about 0.1 ml blood. Contagiously transmittedAmblyomma ticks hang on to iguana hosts for days or weeks. Marineiguanas sleeping alone had 2.0 mobile ticks per night, whileindividuals sleeping in groups had 0.1 to 1.1 mobile ticks pernight. Single iguanas decreased their mobile parasite load to0.2 ticks per night by sleeping on bushes. Experimental nightlytranslocation of iguanas to areas without other sleeping iguanassignificantly increased their mobile parasite burden above levelsencountered by naturally single individuals (n = 4.6 ticks pernight). Creating an experimental group of two animals reducedinfestation with mobile ticks by 59% compared to levels on singleanimals. Over the course of weeks, mobile ectoparasite loadsat grouping sites increased to levels found at single sites,at which point marine iguanas changed sleeping sites. Groupinghad no effect on the prevalence of contagious ticks. Furthermore,grouping did not help to conserve body temperature in Genovesaiguanas, as measured by radiotelemetry. I conclude that marineiguanas group during daytime at microhabitats favored for thermoregulation(predation is absent in this population). Thermoregulation wasnot of prime importance for nightly aggregations, which insteadserved to reduce mobile ectoparasite load. As a minimum costof infestation, I estimate that individuals sleeping alone wouldhave a 5.4% lower annual energy budget due to tissue removal,not including potential internal infections.     

Nitrogen excretion during embryonic development of the green iguana, Iguana iguana (Reptilia; Squamata)

Development within the cleidoic egg of birds and reptiles presents the embryo with the problem of accumulation of wastes from nitrogen metabolism. Ammonia derived from protein catabolism is converted into the less toxic product urea or relatively insoluble uric acid. The pattern of nitrogen excretion of the green iguana, Iguana iguana, was determined during embryonic development using samples from allantoic fluid and from the whole homogenized egg, and in hatchlings and adults using samples of blood plasma. Urea was the major excretory product over the course of embryonic development. It was found in higher concentrations in the allantoic sac, suggesting that there is a mechanism present on the allantoic membrane enabling the concentration of urea. The newly hatched iguana still produced urea while adults produced uric acid. The time course of this shift in the type of nitrogen waste was not determined but the change is likely to be related to the water relations associated with the terrestrial habit of the adult. The green iguana produces parchment-shelled eggs that double in mass during incubation due to water absorption; the eggs also accumulate 0.02 mM of urea, representing 82% of the total measured nitrogenous residues that accumulate inside the allantois. The increase in egg mass and urea concentration became significant after 55 days of incubation then were unchanged until hatching.     

The role of urban greenspaces in fauna conservation: the case of the iguana Ctenosaura similis in the ‘patios’ of León city, Nicaragua

Private urban greenspaces, called ‘patios’ in Latin America, can act as important refuges for wildlife in scattered growing cities of the tropics. We studied the presence and abundance of the black spiny-tailed iguana (Ctenosaura similis) in patios of León city (Nicaragua). Forty patios were structurally characterized and abundances of iguanas were determined through surveys of inhabitants supported by observations of specimens and burrows. Patio area and maximum tree height were variables positively related with presence and abundance of iguanas and presence of burrows. The permeability of fences for iguana movements and the presence of preferred trees for food were also related to presence and abundance of iguanas. Stepwise selection model for abundance of iguanas included number of preferred trees, maximum tree height and permeability of fences. The presence of iguanas was only explained by maximum tree height. Our results show that the indigenous-root types of patios offer the most adequate conditions for the establishment and maintenance of iguana populations among the studied cases. In the light of these results, management practices to preserve the Ctenosaura similis populations in urban patios are suggested.     

Water Relations of Chelonian Eggs and Embryos: Is Wetter Better?

The exchange of water between a chelonian egg and its subterraneanenvironment is influenced by numerous factors, the most importantof which are (1) structure of the calcareous layer of the eggshell,(2) water potential and temperature in the nest, and (3) fractionof the eggshell that actually contacts soil in the nest cavity.Eggs with relatively porous shells tend to absorb large quantitiesof water from cool, moist environments and to lose large amountsof water to warm, dry ones. Net water-exchange by such eggsalso tends to be more favorable (= positive) when soil contactsthe entire eggshell than when a large fraction of the shellis exposed to air trapped inside the nest cavity. In contrast,eggs with relatively impermeable shells usually exchange onlysmall amounts of water with their environment, regardless ofthe physical conditions that prevail inside the nest. The patternof net water-exchange, together with size (and water content)of the freshly laid egg, determines the amount of water thatis available to sustain the embryo. An embryo having accessto a relatively large reserve of water will consume more ofits yolk and grow to larger size before hatching than will anembryo having access to a smaller reserve of water. Large, well-hydratedhatchlings may survive better than small, dehydrated animalsduring the trek overland from nest to water. If so, a cooler,wetter nest will also be a better nest.     

Behavioral and Morphological Adaptations by Galapagos Land Iguanas (Conolophus subcristatus) to Water and Energy Requirements of Eggs and Neonates

We examine the importance of both the hydric environment ofnaturally incubating reptilian eggs and the energetic needsof hatchlings via an investigation of reproduction in Galapagosland iguanas (Conolophus subcristatus). Hatching success ofegg clutches and the size of subsequent hatchlings are bothpositively correlated with the water potential of natural nests,as predicted from previous laboratory experiments. Water potentialsrepresenting optimal incubation environments are available foronly a brief period in nature, and depend upon the same seasonalrainfall as does food abundance for the emerging hatchlings.The temporal placement of the reproductive season of Conolophussubcristatus balances these conflicting needs for water by eggsand energy by hatchlings. Oviposition occurs slightly beforesuitable water potentials are reached and hatchling emergenceoccurs after the peak in food abundance. Morphological adaptations by Conolophus subcristatus to theirprecarious reproductive phenology include greater amounts ofalbumen in their eggs, and greater energy reserves in emergenthatchlings than most other lizard species. These adaptationslessen the severity of an arid environment where water becomesavailable for periods too short to allow both oviposition andhatching to be temporally placed in an optimal manner.     

Diet selection and digestion in Iguana iguana: the importance of age and nutrient requirements

Summary The green iguana, Iguana iguana, is herbivorous throughout life, and depends on a microbial fermentation system in the hindgut to degrade plant fiber. Because the metabolic rates of lizards are proportional to body mass raised to the 0.80 power, hatchling iguanas have 2X, and juveniles 1.4X, greater relative energy requirements (kJxg body mass^-1xday^-1) than full-grown adults. Growing animals also need a higher protein intake, for contruction of body tissues, than do mature animals. This study investigated how growing iguanas achieve a relatively greater nutrient intake than adults. Hatchling and juvenile iguanas do not have higher relative capacities of the digestive tract than mature iguanas, nor do they digest plant materials more effectively. Instead, growing iguanas select diets higher in digestible protein, and digest the same food 1.3X to 2X more rapidly, than adults. Young iguanas may accomplish their shorter food transit times by maintaining higher body temperatures.     

Demography and nesting ecology of green iguana, Iguana iguana (Squamata: Iguanidae), in 2 exploited populations in Depresión Momposina, Colombia

We studied the demography and nesting ecology of two populations of Iguana iguana that face heavy exploitation and habitat modification in the Momposina Depression, Colombia. Lineal transect data was analyzed using the Fourier model to provide estimates of social group densities, which was found to differ both within and among populations (1.05-6.0 groups/ha). Mean group size and overall iguana density estimates varied between populations as well (1.5-13.7 iguanas/ha). The density estimates were far lower than those reported from more protected areas in Panama and Venezuela. Iguana densities were consistently higher in sites located along rivers (2.5 iguanas/group) than in sites along the margin of marshes, probably due to vegetational differences (1.5 iguanas/group). There was no correlation between density estimates and estimates of relative abundance (number of iguanas seen/hour/person) due to differing detectabilities of iguana groups among sites. The adult sex ratio (1:2.5 males:females) agreed well with other reports in the literature based upon observation of adult social groups, and probably results from the polygynous mating system in this species rather than a real demographic skew. Nesting in this population occurs from the end of January through March and hatching occurs between April and May. We monitored 34 nests, which suffered little vertebrate predation, perhaps due to the lack of a complete vertebrate fauna in this densely inhabited area, but nests suffered from inundation, cattle trampling, and infestation by phorid fly larvae. Clutch sizes in these populations were lower than all other published reports except for the iguana population on the highly xeric island of Cura?ao, implying that adult females in our area are unusually small. We argue that this is more likely the result of the exploitation of these populations rather than an adaptive response to environmentally extreme conditions.     

The effect of the thermal environment on the ability of hatchling Galapagos land iguanas to avoid predation during dispersal

Summary During the month of February 1979, several hundred hatchling land iguanas (Conolophus pallidus) were observed emerging from their natal burrows in a 2 ha communal nesting area on Isla Santa Fe, Galapagos Islands. During this emergence, as many as nine Galapagos hawks were observed to patrol the nesting area and attack hatchling iguanas.The hypothesis that the ability of hatchling land iguanas to escape predation could be influenced by the interaction of the physiological state of the lizards and the thermal environment was analyzed using (1) empirical data on the effect of body temperature (T _b) on locomotory ability of iguanas and (2) biophysical modeling of the T _b's of hatchlings under natural conditions. This hypothesis was tested by assessing the success of natural hawk attacks on lizards exposed to different thermal environments.During those periods when predicted T _b's of hatchlings were always <32°C, (at which temperatures land iguanas were shown to have less than maximal ability to sprint rapidly) hawks were successful in 67% of the observed attacks. However, when T _b's of hatchlings were always 32° C, hawks were successful on only 19% of observed attacks. During periods when hatchling T _b's could be <32° C or 32–40° C (depending upon which microhabitat the hatchling occupied before the attack), the hawks were successful in 46% of the observed attacks.These data indicate that the physical environment, as mediated through the physiological state of the lizards and to correlated locomotary abilities, significantly affects the ability of hatchling land iguanas to escape predation.     

Optimal foraging of a herbivorous lizard,the green iguana in a seasonal environment

Food selection was studied in free living green iguanas (Iguana iguana) throughout the year in a semiarid environment, Curaçao (Netherlands Antilles). Food intake was determined by direct observations and converted into biomass intake. Comparison between intake and biomass availability of the various food items revealed that the lizards were selective, and that changes in seasonal food availability led to periodic switching of food plants. The extent to which nutrient constraints determine iguana feeding ecology was investigated. Potential constraints were the requirements for water, digestible crude protein, and metabolizable energy. By using a linear programming model that incorporates characteristics of the food (chemical composition, energy content, item size) and requirements and constraints of the green iguanas (nutrient and energy requirements digestive tract capacity, feeding rate) it was possible to identify which factors determine food choice over the year. During the dry period, when the iguanas had no access to drinking water they consumed flowers to increase water intake, though the amount of flowers consumed was too low to cover maintenance requirements for either energy or protein. After the young leaf flush, following the early rains in May, the biomass increased, free surface water was available during showers, and the linear programming solutions indicate that food selection conformed to the protein maximization criterion. Reproduction in green iguanas shows an annual cycle, in which oviposition takes place at the end of the dry season, when intake is below maintenance levels. Females show a 8–10 month gap between acquisition of most of the protein required for egg synthesis and the act of laying. Thus, as in avian and mammalian herbivores, food availability during a period prior to the energy and protein demanding reproductive season of iguanas determines reproductive success.     

Taphonomy of modern and fossil intertidal gastropod associations from Isla Santa Cruz and Isla Santa Fe, Galápagos Islands

Examination of modern gastropod associations from the low intertidal zone of Isla Santa Cruz suggests that fossil rocky intertidal deposits from this tropical locality will be taphonomically compromised in three ways: (1) Marine hermit crabs, by their use of empty gastropod shells, will mix the shells from varying tidal heights and habitats, thus facilitating mixed associations of such shells in the fossil record, (2) encrusting organisms on crab-inhabited shells are abundant, while boring organisms are almost non-existent, indicating possible differences in postmortem shell retention, and (3) intertidal shells are further taphonomically altered by terrestrial hermit crabs, which transport the shells onto land as well as physically modify the shells. Gastropod fossils from beach and terrace deposits on Isla Santa Fe are interpreted to be a mixed assemblage of rocky intertidal assemblage with few shells indicating influence from marine hermit crabs. Modification of the shell by marine and terrestrial hermit crabs was also evident. A unique polish to the shells at one locality is attributed to the marine iguanas and is only found in the terrace site biologically bulldozed by egg-laying iguanas. Few studies exist on modern rocky intertidal associations in the Galápagos, and the fossil record of rocky shores may provide a baseline for future studies in how community structure has changed over since the advent of humans. Galapagos, C oenobita C ompressus , gastropods, humans, Gulf of California, bionts, nutrients.
Sally E. Walker, Department of Geology, The University of Georgia, Athens, Georgia, USA; 8th September, 1994; revised 28th June, 1995.     

Ty1-copia group retrotransposon sequences in amphibia and reptilia

We have isolated sequences belonging to Tyl-copia group retrotransposons from the genomes of an amphibian (Pyxicephalus adspersa) and three reptiles (Conolophus subscristatus, Amblyrynchus cristatus and Pytas mucosus). Two different seqences were found in the amphibian (Tpa1 and Tpa2). Each is present in several copies per genome and absent from the genomes of two other amphibian species. The C. subcristatus sequence Tcs1 is present in multiple copies in both its host genome (Galapagos land iguana) and the genome of the related Galapagos marine iguana (A. cristatus). There is little or no polymorphism in Tcs1 insertions between different individual animals, suggesting that this sequence is not transposing rapidly in either iguana genome. The P. mucosus sequence Tpm1 shows a discontinuous distribution in snake species, suggesting that it has either been lost from many lineages during vertical germline transmission or has been transferred horizontally in some snake species. Phylogenetic comparisons of all these sequences with each other and with other members of this retrotransposon group from other animals and plants show that sequences within a particular vertebrate species are most closely related to each other, consistent with a vertical transmission model for their evolution.     

Ty1-copia group retrotransposon sequences in amphibia and reptilia

We have isolated sequences belonging to Tyl-copia group retrotransposons from the genomes of an amphibian (Pyxicephalus adspersa) and three reptiles (Conolophus subscristatus, Amblyrynchus cristatus and Pytas mucosus). Two different seqences were found in the amphibian (Tpa1 and Tpa2). Each is present in several copies per genome and absent from the genomes of two other amphibian species. The C. subcristatus sequence Tcs1 is present in multiple copies in both its host genome (Galapagos land iguana) and the genome of the related Galapagos marine iguana (A. cristatus). There is little or no polymorphism in Tcs1 insertions between different individual animals, suggesting that this sequence is not transposing rapidly in either iguana genome. The P. mucosus sequence Tpm1 shows a discontinuous distribution in snake species, suggesting that it has either been lost from many lineages during vertical germline transmission or has been transferred horizontally in some snake species. Phylogenetic comparisons of all these sequences with each other and with other members of this retrotransposon group from other animals and plants show that sequences within a particular vertebrate species are most closely related to each other, consistent with a vertical transmission model for their evolution.     

Predominance of single paternity in the black spiny-tailed iguana: conservation genetic concerns for female-biased hunting

Because of female-biased illegal harvesting, knowledge about the genetic mating system of the black spiny-tailed iguana Ctenosaura pectinata is of primary interest for the conservation of this threatened species. Based on the high levels of multiple paternity found in clutches of many other reptiles, particularly in lizards, it is hypothesised that multiple paternity may also be common in black iguanas. This was investigated by using microsatellite DNA to estimate the number of males siring nine litters (9 mothers, 121 offspring genotyped at ten polymorphic loci) of black iguanas. Contrary to expectations, only 11% of sampled black iguana females produced litters consistent with being sired by multiple males. These data are the first evidence for the predominance of single paternity within an iguanid lizard, and suggest that black iguana may be more susceptible to loss of genetic variation in the face of gender-biased over-hunting pressure than previously thought.     

Seed Structure in Cannaceae: Taxonomic and Ecological Implications

The ovules and seeds of Canna show some striking differencesto those in other zingiberalean families. The pachychalazaldevelopment of the ovule results in a seed of which only a smallpart of the testa is of tegumentary origin. A silicified endotesta,characteristic of the order, is lacking. The mechanical layerof the seed is formed by a continuous exotesta of Malpighiancells. The intact seed coat is impermeable. The seed is ableto absorb water after the raising of a preformed imbibitionlid on the raphe. During imbibition the extotesta loses itshardness, allowing the embryo to emerge. The special structuralfeatures of the seed are discussed in relation to records onthe extreme longevity of Canna seeds. Canna tuerckheimii, C. jaegeriana, C. glauca, pachychalaza, Malpighian cells, imbibition lid, seed longevity