Focusing and accommodation in tuatara (Sphenodon punctatus)

Summary Photorefraction and photokeratometry were performed on two juvenile tuatara (Sphenodon punctatus, 7 years of age, total length approx. 27 cm). Sphenodon is the only surviving genus of the Rhynchocephalia, an order of reptiles. Both existing species are endangered and are found only in New Zealand. Even though Walls (1942) has stated that the ciliary muscle is relatively weak in tuataras, we observed 8 D of accommodation. The eyes were found to focus independently and we could induce an anisometropia of 6 D. The average corneal power of the tuatara was found to be 101 D.     

Morphological changes in the corpus luteum of tuatara (Sphenodon punctatus) during gravidity

Tuatara (Sphenodon spp.) are rare reptiles, members of the reptilian order Sphenodontida, inhabiting small offshore islands of New Zealand. Females usually require about three years to yolk a clutch of eggs followed by an 8-month period of in utero egg shelling. As in other vertebrates, the post-ovulatory follicle forms a transitory endocrine structure, the corpus luteum. The tuatara Sphenodon punctatus exhibits a corpus luteum having several unusual morphological features as compared to turtles and squamate reptiles. Like the crocodilians, the tuatara has a corpus luteum in which the luteal cell mass never fills the central cavity and in which the thecal fibroblasts do not close the ovulation aperture. As in all oviparous reptiles examined, however, the corpus luteum appears to persist throughout gravidity based on its histological appearance. During gravidity, plasma progesterone concentrations are detectable, even though gravidity lasts an exceptionally long time (8 months) for an oviparous species. Luteolysis is initiated within two months following oviposition. The initial stages of luteolysis appear rapid, but luteal scar tissue is apparent in the ovaries of all adult females we examined and probably persists for many years post-oviposition. J Morphol 232:79–91, 1997. © 1997 Wiley-Liss, Inc.     

Eggshell formation during prolonged gravidity of the tuatara Sphenodon punctatus

Scanning electron microscopy (SEM) was used to examine the process of shell formation in tuatara. Tuatara carry eggs in the oviducts for ∼ 7–8 mo before nesting, a period of gravidity more than three times as long as in any other oviparous reptile. Our aim was to determine whether shell formation occurred rapidly after ovulation, or whether it occurred gradually throughout gravidity. Eggs were obtained from females in early gravidity (May, ∼ 1 mo after ovulation), midgravidity (August and September, 4–5 mo after ovulation), and late gravidity, immediately prior to nesting (December, 8 mo after ovulation). The shell membrane (fibrous layer) was well formed by May, but calcification of the outer surface had only just begun. Vertical columns of calcium carbonate were embedded in the shell membrane and appeared to erupt through the outer surface between early and midgravidity. Changes in the appearance of the outer calcareous layer were evident as gravidity progressed. In all shells, calcium carbonate was present as calcite. The appearance of the inner boundary (innermost layer of eggshell) was variable; some shells had a smooth and amorphous inner boundary as previously reported for tuatara and other reptiles, whereas other shells had an inner boundary composed of small spherical granules on the inner surface of which small calcareous spicules were scattered. A previously published model of the process of shell formation in tuatara eggshells is refined in light of our observations. We interpret the ability of female tuatara to shell their eggs gradually during winter as further evidence of their unusual physiological tolerance of cold conditions. © 1996 Wiley-Liss, Inc.     

Mastication in the tuatara,Sphenodon punctatus (reptilia: Rhynchocephalia): Structure and activity of the motor system

The masticatory pattern of Sphenodon punctatus, the sole remaining rhynchocephalian, now restricted to islands off the coast of New Zealand, has been analyzed by detailed anatomy, cinematography, cinefluoroscopy, and electromyography. Food reduction consists of a closing, crushing bite followed by a propalineal sliding of the dentary row between the maxillary and palatine ones. The large, fleshy tongue can be protruded to pick up small prey, and also plays a major role in prey manipulation. The rotational closing movement of the jaw, supporting the basic crushing movement, is induced by the main adductor musculature. It is followed by a propalineal anterior displacement relying heavily on the action of the M. pterygoideus. The fiber lengths of the several muscles reflect the extent of shortening. The most obvious modification appears in the M. pterygoideus, which contains a central slip of pinnately arranged short fibers that act a period different from that of the rest of the muscle; their action increases the power during the terminal portion of the propalineal phase. This also allows the animal to use its short teeth in an effective shearing bite that cuts fragments off large prey. The action of single cusped dentary teeth acting between the maxillary and palatine tooth rows provides a translational crushing-cutting action that may be an analog of the mammalian molar pattern. However, this strictly fore-aft slide does not incorporate capacity for later development of lateral movement.     

Physiological effects of a fish oil supplement on captive juvenile tuatara (Sphenodon punctatus)

Tuatara (Sphenodon, Order Sphenodontia) are rare New Zealand reptiles whose conservation involves captive breeding. Wild tuatara eat seabirds, which contain high levels of the long-chain n-3 polyunsaturated fatty acids (PUFAs) eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA). These fatty acids are absent from the captive diet, and consequently, plasma fatty acid composition of wild and captive tuatara differs. This study investigated the effects of incorporating EPA and DHA into the diet of captive juvenile tuatara (Sphenodon punctatus) in an attempt to replicate the plasma fatty acid composition of wild tuatara. Tuatara receiving a fish oil supplement containing EPA and DHA showed overall changes in their plasma fatty acid composition. Phospholipid EPA and DHA increased markedly, reaching 10.0% and 5.9 mol%, respectively, by 18 mo (cf. 相似文献   

Aspects of shell formation in eggs of the tuatara,Sphenodon punctatus

The flexible shell from eggs of the tuatara (Sphenodon punctatus) is comprised of both calcareous and fibrous components. The calcareous material is organized into columns that extend deep into the fibrous shell membrane. Many of the fibers of the membrane are enclosed within the crystalline matrix of the columns. Columns widen and flatten slightly at the outer surface of the eggshell to form cap-like structures composed of a compact crystalline matrix containing no fibers. The outer surface of eggs laid prior to completion of shell formation consists of a series of nodes obscured by a densely fibrous matrix. Similar nodes also are found at the inner surface of partially shelled eggs. The nodes represent the outer and inner aspects of columns that had not completed formation prior to oviposition. Our interpretation is that a layer (or layers) of the shell membrane forms first, with nucleation of columns occurring shortly thereafter. Columns grow into the membrane a short distance and enclose fibers of the membrane, but the primary direction of column growth is toward what will become the outer aspect of the shell. Calcareous columns and the shell membrane form more or less in concert until crystal growth outstrips that of the membrane and a cap-like apex of compact crystalline material is formed. The end result is an eggshell in which the shell membrane and calcareous material form a single unit for much of the thickness of the shell.     

Distribution and characterization of keratins in the epidermis of the tuatara (Sphenodon punctatus; Lepidosauria, Reptilia)

Reptilian scales are mainly composed of alpha-and beta-keratins. Epidermis and molts from adult individuals of an ancient reptilian species, the tuatara (Sphenodon punctatus), were analysed by immunocytochemistry, mono- and bi-dimensional electrophoresis, and western blotting for alpha- and beta-keratins. The epidermis of this reptilian species with primitive anatomical traits should represent one of the more ancient amniotic epidermises available. Soft keratins (AE1- and AE3-positive) of 40-63 kDa and with isoelectric points (pI) at 4.0-6.8 were found in molts. The AE3 antibody was diffusely localised over the tonofilaments of keratinocytes. The lack of basic cytokeratins may be due to keratin alteration in molts, following corneification or enzymatic degradation of keratins. Hard (beta-) keratins of 16-18 kDa and pI at 6.8, 8.0, and 9.2 were identified using a beta-1 antibody produced against chick scale beta-keratin. The antibody also labeled filaments of beta-cells and of the mature, compact beta-layer. We have shown that beta-keratins in the tuatara resemble those of lizards and snakes, and that they are mainly basic proteins. These proteins replace cytokeratins in the pre-corneoum beta-layers, from which a hard, mechanically resistant corneoum layer is formed over scales. Beta-keratins may have both a fibrous and a matrix role in forming the hard texture of corneoum scales in this ancient species, as well as in more recently evolved reptiles.     

Formation of the corneous layer in the epidermis of the tuatara (Sphenodon punctatus, Sphenodontida, Lepidosauria, Reptilia)

The formation of the stratum corneum in the epidermis of the reptile Sphenodon punctatus has been studied by histochemical, immunohistochemical, and ultrastructural methods. Sulfhydryl groups are present in the mesos and pre-alpha-layer but disappear in the keratinized beta-layer and in most of the mature alpha-layer. This suggests a complete cross-linking of keratin filaments. Tyrosine increases in keratinized layers, especially in the beta-layer. Arginine is present in living epidermal layers, in the presumptive alpha-layer, but decreases in keratinized layers. Histidine is present in corneous layers, especially in the intermediate region between the alpha- and a new beta-layer, but disappears in living layers. It is unknown whether histidine-rich proteins are produced in the intermediate region. Small keratohyalin-like granules are incorporated in the intermediate region. The plane of shedding, as confirmed from the study on molts, is located along the basalmost part of the alpha-layer and may involve the degradation of whole cells or cell junctions of the intermediate region. A specific shedding complex, like that of lizards and snakes, is not formed in tuatara epidermis. AE1-, AE2-, or AE3-positive alpha-keratins are present in different epidermal layers with a pattern similar to that previously described in reptiles. The AE1 antibody stains the basal and, less intensely, the first suprabasal layers. Pre-keratinized, alpha- and beta-layers, and the intermediate region remain unlabeled. The AE2 antibody stains suprabasal and forming alpha- and beta-layers, but does not stain the basal and suprabasal layers. In the mature beta-layer the immunostaining disappears. The AE3 antibody stains all epidermal layers but disappears in alpha- and beta-layers. Immunolocalization for chick scale beta-keratins labels the forming and mature beta-layer, but disappears in the mesos and alpha-layer. This suggests the presence of common epitopes in avian and reptilian beta-keratins. Low molecular weight alpha-keratins present in the basal layer are probably replaced by keratins of higher molecular weight in keratinizing layers (AE2-positive). This keratin pattern was probably established since the beginning of land adaptation in amniotes.     

Observations on the activity and thermoregulation of the tuatara,Sphenodon punctatus (Reptilia: Rhynchocephalia), on Stephens Island

Abstract

Observations on Stephens Island, Cook Strait, in December 1978 show that although the tuatara is generally most active at night, many animals spend much of the day at or beyond burrow entrances, apparently to increase their body temperature. During the day, tuataras tend to move further from burrows which are under shaded forest than from those in open pasture. By day, mean body temperatures (±SE) ranged from 17.2±0.5°C in forest shade to 24.6±1.1°C in full sunlight; the maximum body temperature recorded was 26.3°C. The significance of these observations is discussed.     

Comparative fine structure of the axial skeleton inside the regenerated tail of some lizard species and the tuatara (Sphenodon punctatus)

The regenerated tail of the New Zealand gecko Hoplodactylus maculatus is equipped with an elastic cartilaginous tube as skeletal axis. Other lizard species and Sphenodon punctatus possess variably developed hyaline cartilaginous tubes. Moreover, H. maculatus enhances the functional performance of its tail by long elastic fibres, which are arranged all around the central regenerated spinal cord. The different characteristics of the regenerated skeleton could be related to the different environments that the species studied occupy in nature.     

Morphological,cytochemical, and biochemical observations on the blood of the tuatara,Sphenodon punctatus

Abstract

Morphological features of the circulating blood cells of an adult female tuatara are described and illustrated. Mitosis in circulating erythrocytes and the occurrence of an intraerythrocytic haemogregarine parasite are noted. The packed cell volume (haematocrit) was 35%. There were about 310 000 erythrocytes and 7200 leucocytes per mm³ of blood. The lymphocyte was the predominant leucocyte, followed by the eosinophil, monocyte, neutrophil, and basophil. The eosinophils and neutrophils reacted positively to acid phosphatase and peroxidase; the neutrophils were also esterase positive, and all three granulocytes were PAS positive. Haemoglobin, plasma protein, glucose, sodium, potassium, cholesterol, and triglyceride levels were determined. The primitive nature of the tuatara is reflected in the morphological. mensural, and biochemical data presented.     

Differences in dietary and plasma fatty acids between wild and captive populations of a rare reptile, the tuatara (Sphenodon punctatus)

Tuatara (Sphenodon) are rare reptiles endemic to New Zealand. Wild tuatara on Stephens Island (study population) prey on insects as well as the eggs and chicks of a small nesting seabird, the fairy prion (Pachyptila turtur). Tuatara in captivity (zoos) are fed diets containing different insects and lacking seabirds. We compared the fatty acid composition of major dietary items and plasma of wild and captive tuatara. Fairy prions (eaten by tuatara in the wild) were rich in C20 and C22 polyunsaturated fatty acids (PUFA), especially the n-3 fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA). In contrast, items from the diet of captive tuatara contained no C20 and C22 PUFA and were higher in medium-chain and less unsaturated fatty acids. Plasma from wild tuatara was higher in n-3 PUFA [including alpha-linoleic acid (C18:3n-3), EPA and DHA], and generally lower in oleic acid (C18:1) and palmitic acid (C16:0), than plasma from captive tuatara in the various fractions (phospholipid, triacylglycerol, cholesterol ester and free fatty acids). Plasma from wild adult tuatara showed strong seasonal variation in fatty acid composition, reflecting seasonal consumption of fairy prions. Differences in the composition of diets and plasma between wild and captive tuatara may have consequences for growth and reproduction in captivity. Accepted: 3 August 1998     

Two patterns of variation among MHC class I loci in Tuatara (Sphenodon punctatus)

The genes of the major histocompatibility complex (MHC) are a central component of the immune system in vertebrates and have become important markers of functional, fitness-related genetic variation. We have investigated the evolutionary processes that generate diversity at MHC class I genes in a large population of an archaic reptile species, the tuatara (Sphenodon punctatus), found on Stephens Island, Cook Strait, New Zealand. We identified at least 2 highly polymorphic (UA type) loci and one locus (UZ) exhibiting low polymorphism. The UZ locus is characterized by low nucleotide diversity and weak balancing selection and may be either a nonclassical class I gene or a pseudogene. In contrast, the UA-type alleles have high nucleotide diversity and show evidence of balancing selection at putative peptide-binding sites. Twenty-one different UA-type genotypes were identified among 26 individuals, suggesting that the Stephens Island population has high levels of MHC class I variation. UA-type allelic diversity is generated by a mixture of point mutation and gene conversion. As has been found in birds and fish, gene conversion obscures the genealogical relationships among alleles and prevents the assignment of alleles to loci. Our results suggest that the molecular mechanisms that underpin MHC evolution in nonmammals make locus-specific amplification impossible in some species.     

Status of the tuatara,Sphenodon punctatus,on Hongiora and Ruamahua-iti Islands,Aldermen Group,New Zealand

Abstract

Results from a survey of the tuatara populations on Hongiora and Ruamahua-iti Islands, conducted in March 1986, do not support an earlier report that the Hongiora population is declining (Crook 1973). On Hongiora, 49 captures were made of 43 tuatara over 12 man-hours, and on Ruamahua-iti, 67 captures of 60 tuatara were made over 12.5 man-hours. The mean body lengths of Hongiora tuatara (male=250.4 mm, female=220.8 mm) were significantly larger than those of Ruamahua-iti animals (male= 187.6mm, female= 179.3 mm). We question the interpretation that the smaller average size of tuatara on Ruamahua-iti Island indicates that they are younger than those on Hongiora Island.     

Conservation implications of a long-term decline in body condition of the Brothers Island tuatara (Sphenodon guntheri)

Understanding the factors that drive the dynamics of remnant populations of long-lived species presents a unique challenge for conservation management. The long-lived Brothers Island tuatara Sphenodon guntheri is represented by one natural, self-sustaining population on 4-ha North Brother Island, New Zealand, and two small, translocated populations. The North Brother Island population was almost driven to extinction by extreme habitat modification and collecting in the late 19th century. Analysis of a long-term (1957–2001) dataset, following the population's recovery, reveals a significant decline in tuatara body condition over time, which is more pronounced in females. Declining body condition, coupled with very low reproductive output, may be symptomatic of a density-dependent response to elevated population size exacerbated by resource limitation. Sex-specific effects that disadvantage females could compromise this small population, particularly as it exhibits a male-biased sex ratio. We recommend removal of infrequently used structures and habitat restoration to alleviate intense resource competition. Population-level manipulation should be considered if future monitoring indicates an increasingly male-biased sex ratio and continued decline of female body condition.