Sphenodon guntheriBrothers Island tuatara

Ge­o­graphic Range

Broth­ers Is­land tu­ataras in­habit North Brother Is­land in Cook Straight, New Zealand (41°06′S, 174°26′E). Only a few hun­dred in­di­vid­u­als re­main on North Brother Is­land. (Cree, 1994; Lutz, 2005; Mitchell, et al., 2008; Thomp­son, et al., 1992)

Habi­tat

Broth­ers Is­land tu­ataras are one of the few rep­tiles with the abil­ity to thrive in cooler con­di­tions. They are noc­tur­nal and phys­i­cally ca­pa­ble of with­stand­ing tem­per­a­tures as low as 9°C, with hu­mid­ity in the range of 70 to 80%. High hu­mid­ity and low tem­per­a­tures allow tu­ataras to main­tain healthy shed cy­cles and live longer life spans, due to their ef­fects on heart and meta­bolic rates. Dur­ing the day, most in­di­vid­u­als in­habit bur­rows along cliff faces. Their bur­rows can mea­sure about 5 me­ters in length and 30 cen­time­ters in depth, and are some­times taken over from pre­vi­ous in­hab­i­tants. Bur­rows are typ­i­cally found in open areas fea­tur­ing low coastal veg­e­ta­tion, and usu­ally offer both shade and sun­light to aid in heat reg­u­la­tion. Cer­tain cliffs and other areas of the is­land that pro­vide dif­fer­ent types of ter­rain are often in­hab­ited by birds or other an­i­mals that can com­pete with tu­ataras for ter­ri­tory. (Cree, 1994; Lutz, 2005; Mitchell, et al., 2008; Nel­son, et al., 2002a; Ram­stad, et al., 2007)

  • Range elevation
    87 (high) m
    285.43 (high) ft

Phys­i­cal De­scrip­tion

Tu­ataras gen­er­ally have a lizard-like ap­pear­ance, but dif­fer from lizards in that their teeth are at­tached to bone, they have two tem­po­ral open­ings, they have no ex­ter­nal ear, and the males lack sex­ual or­gans. Tu­atara means “bear­ing spines”, re­fer­ring to the sin­gle row of spines run­ning along their dor­sal side. The skin of this species is gen­er­ally olive-brown with yel­low­ish patches, which of­fers ef­fec­tive cam­ou­flage in their en­vi­ron­ment. When born, tu­ataras pos­sess a third eye on the top of their head (called a pari­etal eye). This pineal spot ap­pears some­what func­tional at birth, but be­comes cov­ered with skin after sev­eral months and does not ap­pear to serve a func­tional pur­pose there­after. Adults are fairly large and rather slow mov­ing, reach­ing a weight of 900 g and a length of 76 cm. Males are larger than fe­males and have pro­por­tion­ately larger heads and crests. (Cree, 1994; Lutz, 2005; Pope, 1956)

Broth­ers Is­land tu­ataras be­long to the order Ryn­cho­cephalia, which con­tains only one other liv­ing species, Sphen­odon punc­ta­tus (spot­ted tu­ataras). Broth­ers Is­land tu­ataras are char­ac­ter­is­ti­cally smaller and have longer re­pro­duc­tive cy­cles than spot­ted tu­ataras. (Cree, 1994; Lutz, 2005; Pope, 1956)

  • Sexual Dimorphism
  • male larger
  • Range mass
    900 (high) g
    31.72 (high) oz
  • Range length
    76 (high) cm
    29.92 (high) in

De­vel­op­ment

In tu­ataras, in­cu­ba­tion tem­per­a­tures above 22°C tend to pro­duce males, while tem­per­a­tures of 20°C and below re­sult in more fe­male off­spring. Young tu­ataras es­cape their eggs by using an egg tooth. This struc­ture is lo­cated on the tip of their head be­tween the nos­trils and is lost after the first cou­ple of weeks. Newly hatched tu­ataras re­sem­ble minia­ture ver­sions of adults, and grow very slowly, tak­ing as long as 35 years to reach adult sizes. (Cree, 2002; Lutz, 2005; Mitchell, et al., 2006; Nel­son, et al., 2002a; Nel­son, et al., 2010)

Re­pro­duc­tion

Lit­tle is known about the so­cial struc­ture of mat­ing sys­tems in tu­ataras, but males tend to be highly ter­ri­to­r­ial and mate with mul­ti­ple fe­males if given the chance. Male tu­ataras gen­er­ally out­num­ber fe­males in their na­tive en­vi­ron­ments. Cop­u­la­tion con­sists of a male mount­ing a fe­male and ex­cret­ing sperm from the cloaca. (Cree, et al., 1991; Cree, 1994; Lutz, 2005; New­man, et al., 1994)

Fe­male tu­ataras reach sex­ual ma­tu­rity at 10 to 20 years. Tu­ataras have long life spans and pro­longed re­pro­duc­tive cy­cles. Fe­males usu­ally dig nests in soil lo­cated on cliff edges. Tu­ataras on North Brother Is­land pro­duce an av­er­age of 1.27 eggs per year for each ma­ture fe­male. The mean clutch size of Broth­ers Is­land tu­ataras is ap­prox­i­mately 6.5 eggs. Each egg has a mean weight of 4.9 grams, and the shell has a white col­oration with a rather soft tex­ture. Tu­ataras may lay eggs as often as every 2 years, but most lay eggs every 4 to 5 years. They mate in late sum­mer (De­cem­ber through Feb­ru­ary in New Zealand), with eggs being layed the fol­low­ing spring. (Cree, et al., 1991; Cree, 1994; Hall, 2007; Hemphill, 2012; New­man, et al., 1994)

  • Breeding interval
    Tuataras typically breed every 4 to 5 years.
  • Breeding season
    Tuataras mate in late summer and lay eggs during the spring. Eggs undergo an incubation period of 12-16 months.
  • Average number of offspring
    6.5
  • Range gestation period
    8 to 10 months
  • Range age at sexual or reproductive maturity (female)
    10 to 20 years

Fe­males in­vest en­ergy in their young via the pro­duc­tion of egg yolks and shells. Once the eggs are laid, nei­ther par­ent pro­tects the eggs. There is also no parental in­vest­ment post-hatch­ing. (Hall, 2007)

  • Parental Investment
  • no parental involvement

Lifes­pan/Longevity

There is some de­bate re­gard­ing the full ex­tent of tu­atara lifes­pans, but they are known to be able to live for over 100 years. Their longevity is mainly due to their slow me­tab­o­lism and low body tem­per­a­tures. Lit­tle is known about the lifes­pan of tu­ataras in cap­tiv­ity, as they are not gen­er­ally kept as pets. (Lutz, 2005; Nel­son, et al., 2002b; Ram­stad, et al., 2007; Thomp­son, et al., 1992)

  • Range lifespan
    Status: wild
    100 (high) years
  • Typical lifespan
    Status: wild
    60 to 100 years

Be­hav­ior

The weather plays a role in tu­atara ac­tiv­ity lev­els. Even though they are ca­pa­ble of with­stand­ing lower tem­per­a­tures than most other rep­tiles (as low as 9°C), their ac­tiv­ity level will de­cline in colder weather. There is some de­bate as to whether tu­ataras ac­tu­ally hi­ber­nate, but it is be­lieved that on colder nights they can main­tain a sort of tor­por. Hu­mid­ity on North Brother Is­land is typ­i­cally 70-89%; lower per­cent­ages can re­sult in ac­tiv­ity sup­pres­sion and a pref­er­ence to re­main in bur­rows. (Hall, 2007; Lutz, 2005; Walls, 1983)

Older tu­ataras are ac­tive from dusk until dawn and emerge from bur­rows mostly at night. Young tu­ataras are often di­ur­nal, as their faster speed al­lows them to es­cape pre­da­tion. Older tu­ataras are will set up ter­ri­to­ries that they de­fend with dis­plays or phys­i­cal at­tacks. The most com­monly per­formed ter­ri­to­r­ial dis­plays in­clude for­ward ap­proaches, in­fla­tion of the body, rais­ing of the crest, head bob­bing from side to side, and mouth gap­ing. (Gilling­ham, et al., 1995; Tere­zow, et al., 2008)

  • Range territory size
    11.5 to 86.7 m^2

Home Range

De­spite being ter­ri­to­r­ial, tu­ataras typ­i­cally live in close prox­im­ity to each other. Males be­come ag­gres­sive when other male tu­ataras enter their ter­ri­tory. Fe­males will also be­come ag­gres­sive to­wards other fe­males, but often per­mit males into their ter­ri­tory. Ter­ri­tory sizes are 11.5 to 86.7 m^2. Fe­male ter­ri­to­ries are typ­i­cally half the size of males' and in some cases over­lap­ping ter­ri­to­ries can occur be­tween sexes. (Gilling­ham, et al., 1995; Hall, 2007; Lutz, 2005; Tere­zow, et al., 2008; Walls, 1983)

Com­mu­ni­ca­tion and Per­cep­tion

The pupils of the tu­atara read­ily ex­pand and con­tract to help them see di­ur­nally and at night. Al­though they have no ex­ter­nal ears, they are still able to hear. They are also able to use touch, smell, and taste to per­ceive their en­vi­ron­ment. (Lutz, 2005; Naskrecki, 2011; O'Shea and Hal­l­i­day, 2002)

Tu­ataras be­come ter­ri­to­r­ial at about 6 months of age. Males often in­flate their bod­ies, chase off ri­vals, head bob, gape their mouth, and raise their crests in order to de­fend their ter­ri­to­ries. Dur­ing breed­ing sea­son males may croak, which is used as a mat­ing call to alert fe­males to their pres­ence. (Gilling­ham, et al., 1995; Lutz, 2005)

Food Habits

Tu­ataras are car­niv­o­rous and will eat what­ever they can catch. They often prey on bee­tles, worms, lizards, and other tu­ataras. They pre­fer to eat wetas (Dein­crida ru­gosa), an in­sect species that is en­demic to New Zealand. Oc­ca­sion­ally, tu­ataras will eat sea bird eggs lo­cated in bor­rows close to their ter­ri­to­ries. (Hall, 2007; Mitchell, et al., 2008; Naskrecki, 2011; New, 2008)

  • Animal Foods
  • birds
  • reptiles
  • eggs
  • insects
  • terrestrial worms

Pre­da­tion

Sea bird species that in­habit North Brother Is­land some­times at­tack tu­ataras, often for ter­ri­to­r­ial rea­sons. How­ever, swamp har­ri­ers (Cir­cus ap­prox­i­mans) and New Zealand fal­cons (Falco no­vaezee­landiae) are known to catch and con­sume younger tu­ataras. In­va­sive species like rats also pre­date on tu­ataras. Young tu­ataras are di­ur­nal, which, along with their faster speed, helps them avoid being prey to older mem­bers of their species. Tu­ataras may also drop and re­gen­er­ate their tails in order to es­cape pre­da­tion. (Ching, 1986; Lutz, 2005; Meads, 1990; Selig­mann, et al., 2008; Tere­zow, et al., 2008)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

Tu­ataras prey mostly on in­ver­te­brates. As tu­ataras and sea birds may live in close prox­im­ity, tu­ataras oc­ca­sion­ally steal eggs from the birds. The main preda­tors of tu­ataras in­clude birds (Falco no­vaezee­landiae and Cir­cus ap­prox­i­mans), dogs, and rats. A species of tick (Am­bly­omma sphen­odonti) has been doc­u­mented as an ex­ter­nal par­a­site of this species. As the spe­cific ep­i­thet of this tick in­di­cates, it is only found on tu­ataras. (Lutz, 2005; Ram­stad, et al., 2007; Tere­zow, et al., 2008)

Com­men­sal/Par­a­sitic Species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Tu­ataras' un­usual phys­i­cal char­ac­ter­is­tics and low pop­u­la­tion num­bers at­tract at­ten­tion and fund­ing from sci­en­tists and con­ser­va­tion­ists. How­ever, there are no known pos­i­tive ben­e­fits of tu­ataras to na­tives of the is­lands where they are found. Al­though tu­ataras are il­le­gal in the pet trade, they have been placed on the black mar­ket for thou­sands of U.S. dol­lars. (Lutz, 2005; O'Shea and Hal­l­i­day, 2002; Ram­stad, et al., 2007)

  • Positive Impacts
  • research and education

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known neg­a­tive eco­nomic ef­fects of Broth­ers Is­land tu­ataras on hu­mans. When threat­ened or han­dled, they are ca­pa­ble of de­liv­er­ing a painful bite. (Lutz, 2005; Ram­stad, et al., 2007)

  • Negative Impacts
  • injures humans
    • bites or stings

Con­ser­va­tion Sta­tus

Human de­vel­op­ment and in­tro­duc­tion of in­va­sive species on North Brother Is­land ap­pears to have caused the most dam­age to pop­u­la­tion num­bers in tu­ataras. Even though pop­u­la­tion num­bers are low and Broth­ers Is­land tu­ataras ex­hibit a male-ori­ented sex ratio, long-term sur­vival is prob­a­ble. Tu­ataras do not re­pro­duce often, al­though their long life spans helps with their over­all con­ser­va­tion. (Cree, et al., 1991; Hoare, et al., 2006; Lutz, 2005; Nel­son, et al., 2002a; Ram­stad, et al., 2007)

Broth­ers Is­land tu­ataras are pro­tected by the gov­ern­ment of New Zealand. It is un­law­ful to col­lect tu­ataras for pets or kill them for any rea­son. Due to low pop­u­la­tion num­bers, many or­ga­ni­za­tions have rec­og­nized that sci­en­tific re­search on cap­tive tu­ataras is nec­es­sary to con­serve this species and keep their ge­netic di­ver­sity as high as pos­si­ble. Vic­to­ria Uni­ver­sity is ac­tively in­volved in the long-term sur­vival of the tu­atara through stud­ies of cap­tive tu­ataras in­di­vid­u­als. ("Tu­atara Re­cov­ery Plan 2001-2011", 2001; Lutz, 2005)

Translo­ca­tion of Broth­ers Is­land tu­ataras has been at­tempted. How­ever, there is much de­bate as to whether this is the most use­ful method for in­creas­ing their pop­u­la­tion size. Nu­tri­ents and ter­ri­tory on North Brother Is­land are lim­ited and too many tu­ataras might be re­lo­cated in the same place. This could po­ten­tially harm na­tive and new tu­atara pop­u­la­tions. If food and ter­ri­tory are not lim­it­ing, translo­cated tu­ataras ap­pear to adapt well to their new en­vi­ron­ment. ("Tu­atara Re­cov­ery Plan 2001-2011", 2001; Nel­son, et al., 2002b)

Other Com­ments

A re­cent mol­e­c­u­lar study uti­liz­ing mi­crosatel­lite and mi­to­chon­dr­ial DNA con­cluded that the genus Sphen­odon con­tains only one species, and that Broth­ers Is­land tu­ataras rep­re­sent a ge­o­graphic vari­ant of this more broadly dis­trib­uted species. De­spite this study, many re­searchers and gov­ern­ment agen­cies con­tinue to rec­og­nize Broth­ers Is­land tu­ataras as a dis­tinct species. (Hay, et al., 2010)

The Maori tribe have few leg­ends based on the Broth­ers Is­land tu­atara. Of the peo­ple that sub­scribe to these leg­ends, some be­lieve tu­ataras are bless­ings while oth­ers be­lieve tu­ataras act as omens. (Naskrecki, 2011; Ram­stad, et al., 2007)

Con­trib­u­tors

Amanda Johns (au­thor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor), Rad­ford Uni­ver­sity, Kier­sten Newtoff (ed­i­tor), Rad­ford Uni­ver­sity, Melissa Whistle­man (ed­i­tor), Rad­ford Uni­ver­sity, Je­remy Wright (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

indeterminate growth

Animals with indeterminate growth continue to grow throughout their lives.

insectivore

An animal that eats mainly insects or spiders.

island endemic

animals that live only on an island or set of islands.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

polygynous

having more than one female as a mate at one time

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

visual

uses sight to communicate

Ref­er­ences

New Zealand De­part­ment of Con­ser­va­tion. Tu­atara Re­cov­ery Plan 2001-2011. Threat­ened Species Re­cov­ery Plan 47. Welling­ton, New Zealand: New Zealand De­part­ment of Con­ser­va­tion. 2001.

Ching, R. 1986. New Zealand Birds: An Artist's Field Stud­ies. Cor­nell Uni­ver­sity: Reed Methuen.

Cree, A. 1994. Low an­nual re­pro­duc­tive out­put in fe­male rep­tiles from New Zealand. New Zealand Jour­nal of Zo­ol­ogy, 21/4: 351-372.

Cree, A. 2002. Tu­atara. Pp. 210-211 in T Hal­l­i­day, K Adler, eds. The New En­cy­clo­pe­dia of Rep­tiles and Am­phib­ians. Ox­ford, UK: Ox­ford Uni­ver­sity Press.

Cree, A., C. Daugh­erty, S. Schafer, D. Brown. 1991. Nest­ing and clutch size of tu­atara (Sphen­odon gun­theri) on North Brother Is­land, Cook Strait. Tu­atara, 31/1: 9-16.

Gilling­ham, J., C. Carmichael, T. Miller. 1995. So­cial be­hav­ior of the tu­atara, Sphen­odon punc­ta­tus. Her­peto­log­i­cal Mono­graphs, 9/1: 5-16.

Hall, D. 2007. The Ul­ti­mate Guide to Snakes and Rep­tiles. Edi­son, New Jer­sey: Chartwell Books, Inc.

Hay, J., S. Sarre, D. Lam­bert, F. Al­len­dorf, C. Daugh­erty. 2010. Ge­netic di­ver­sity and tax­on­omy: a re­assess­ment of species des­ig­na­tion in tu­atara (Sphen­odon: Rep­tilia). Con­ser­va­tion Ge­net­ics, 11: 1063-1081.

Hemphill, K. 2012. DK Eye­wit­ness Travel Guide: New Zealand. Pen­guin.

Hoare, J., S. Pledger, S. Keall, N. Nel­son, N. Mitchell, C. Daugh­erty. 2006. Con­ser­va­tion im­pli­ca­tions of a long-term de­cline in body con­di­tion of the Broth­ers Is­land tu­atara (Sphen­odon gun­theri). An­i­mal Con­ser­va­tion, 9/4: 456-462.

Lutz, D. 2005. Tu­atara: A Liv­ing Fos­sil. Salem, OR: DIMI Press.

Meads, M. 1990. For­got­ten Fauna: The Rare, En­dan­gered, and Pro­tected In­ver­te­brates of New Zealand. Uni­ver­sity of Cal­i­for­nia: DSIR Pub.

Mitchell, N., M. Kear­ney, N. Nel­son, W. Porter. 2008. Pre­dict­ing the fate of a liv­ing fos­sil: how will global warm­ing af­fect sex de­ter­mi­na­tion and hatch­ing phe­nol­ogy in tu­atara?. The Royal So­ci­ety, 275/1648: 2185-2193.

Mitchell, N., N. Nel­son, A. Cree, S. Pledger, S. Keall, C. Daugh­erty. 2006. Sup­port for a rare pat­tern of tem­per­a­ture-de­pen­dent sex de­ter­mi­na­tion in ar­chaic rep­tiles: ev­i­dence from two species of tu­atara (Sphen­odon). Fron­tiers in Bi­ol­ogy, 3/9: doi:10.1186/1742-9994-3-9.

Naskrecki, P. 2011. Relics: Trav­els in Na­ture's Time Ma­chine. Chicago and Lon­don: The Uni­ver­sity of Chicago Press.

Nel­son, N., S. Keall, D. Brown, C. Daugh­erty. 2002. Es­tab­lish­ing a new wild pop­u­la­tion of tu­atara (Sphen­odon gun­theri). Con­ser­va­tion Bi­ol­ogy, 16/4: 887-894.

Nel­son, N., S. Keall, S. Pledger, C. Daugh­erty. 2002. Male-bi­ased sex ratio in a small tu­atara pop­u­la­tion. Jour­nal of Bio­geog­ra­phy, 29/5-6: 633-640.

Nel­son, N., M. Thomp­son, S. Pledger, S. Keall, C. Daugh­erty. 2006. Per­for­mance of ju­ve­nile tu­atara de­pends on age, clutch, and in­cu­ba­tion regime. Jour­nal of Her­petol­ogy, 40/3: 399-403.

Nel­son, N., J. Moore, S. Pil­lai, S. Keall. 2010. Ther­mosen­si­tive pe­riod for sex de­ter­mi­na­tion in the tu­atara. Sym­po­sium: Rep­tile Re­pro­duc­tion, 5/2: 324-329.

New, T. 2008. In­sect Con­ser­va­tion and Is­lands. Springer.

New­man, D., P. Wat­son, I. Mc­Fad­den. 1994. Egg pro­duc­tion by tu­atara on Lady Alice and Stephens Is­land, New Zealand. New Zealand Jour­nal of Zo­ol­ogy, 21/4: 387-398.

O'Shea, M., T. Hal­l­i­day. 2002. Rep­tiles and Am­phib­ians. Penn­syl­va­nia State Uni­ver­sity: Dk Pub.

Pope, C. 1956. The Rep­tile World. Lon­don: Rout­ledge & Kegan Paul Ltd.

Ram­stad, K., N. Nel­son, G. Paine, D. Beech, A. Paul, P. Paul, F. Al­len­dorf, C. Daugh­tery. 2007. Species and cul­tural con­ser­va­tion in New Zealand: Maori tra­di­tional eco­log­i­cal knowl­edge of tu­atara. Con­ser­va­tion Bi­ol­ogy, 21/2: 455-464.

Selig­mann, H., J. Moravec, Y. Werner. 2008. Mor­pho­log­i­cal, func­tional and evo­lu­tion­ary as­pects of tail au­to­tomy and re­gen­er­a­tion in the ‘liv­ing fos­sil’ Sphen­odon (Rep­tilia: Rhyn­cho­cephalia). Bi­o­log­i­cal Jour­nal of the Lin­nean So­ci­ety, 93/4: 721-743.

Tere­zow, M., N. Nel­son, T. Mark­well. 2008. Cir­ca­dian emer­gence and move­ment of cap­tive ju­ve­nile tu­atara (Sphen­odon spp.). New Zealand Jour­nal of Zo­ol­ogy, 35/3: 205-216.

Thomp­son, M., C. Daugh­tery, A. Cree, D. French, J. Gilling­ham, R. Bar­wick. 1992. Sta­tus and longevity of the tu­atara, Sphen­odon gun­theri, and Du­vau­cel's gecko, Ho­plo­dacty­lus du­vaucelii, on North Brother Is­land, New Zealand. Jour­nal of the Royal So­ci­ety of New Zealand, 22/2: 123-130.

Walls, G. 1983. Ac­tiv­ity of the tu­atara and its re­la­tion­ships to weather con­di­tions on Stephens Is­land, Cook Strait, with ob­ser­va­tions on geckos and in­ver­te­brates. New Zealand Jour­nal of Zo­ol­ogy, 10/3: 309-318.