Incubation behaviour and egg physiology of kiwi (

Incubation behaviour varies among the different taxa of kiwi. For North Island brown kiwi (Apteryx mantelli) and little spotted kiwi (A. owenii), only the male incubates the eggs, except for in the first week. Meanwhile, for Okarito brown kiwi (A. mantelli) and the tokoeka (A. australis), incubation is shared by both sexes. In addition, amongst southern tokoeka, family group members can assist with incubation to the extent that breeding males may take no part in incubation at all. This study shows that kiwi turn their eggs regularly, an observation that contradicts earlier findings. There is a mean incubation temperature of 36.5°C at the top of the egg but the bottom may be 10°C lower in North Island brown kiwi and 5-8°C lower in Okarito brown kiwi eggs. Experiments inducing water loss from fertile and infertile eggs show that the presence of an embryo does not influence rate of water loss. North Island brown kiwi, Okarito brown kiwi and tokoeka regularly have more than one clutch in a season, and some North Island brown kiwi females lay up to seven eggs each year.     

Role of predation in the decline of kiwi,

Kiwi have declined markedly in abundance and range since human settlement of New Zealand. Three of the four species are still extant in mainland forests, despite decades of co-existence with various introduced mammals. Little spotted kiwi is now probably confined to offshore islands. The role of introduced mammals in these population declines was evaluated by measuring the survival rates of adults, eggs and chicks of brown kiwi (A. mantelli) and great spotted kiwi (A. haastii) in mainland forests. Mortality rates of adults ranged from 5%-16% and did not differ significantly between species or sexes. Overall, 14 out of 209 adult kiwi died during 159.6 radio-tracking years. Predators definitely caused five of these deaths. Sixty-nine (68%) of 102 eggs from 77 nesting attempts by 48 pairs failed to hatch. Predators probably caused about 10% of egg failures. Only three of 49 chicks probably survived to adulthood, indicating a juvenile mortality rate of about 94%. Predators killed at least 8% of chicks, 45% of juveniles, and possibly as many as 60% of all young kiwi. Ferrets and dogs were the main predators of adult kiwi, possums and mustelids were the main egg predators, while stoats and cats were largely responsible for the deaths of young kiwi. Population models show that northern brown kiwi are currently declining at 5.8% per annum. This decline could be halted by cutting the current predation rates on young kiwi by about 34% to 33%.     

The breeding season of three species of kiwi (Apteryx) in captivity as determined from egg-laying dates

Records from kiwi ( Apteryx spp.) breeding centres were used to obtain laying dates of 403 North Island brown kiwi ( Apteryx australis mantelli ) eggs, 24 great spotted kiwi ( Apteryx haastii ) eggs and 25 little spotted kiwi ( Apteryx owenii ) eggs. North Island brown kiwi outdoors had an annual cycle of egg-laying with 88.5% of eggs laid from June to January. Eggs were laid in every month of the year, with fewest eggs in April (2.1 % of all eggs). The greatest number of eggs was laid in July (15.6% of all eggs), with a second peak of laying in October (13.5% of all eggs). Both the great spotted kiwi and little spotted kiwi had annual cycles of egg-laying, with the main egg-laying seasons being August-January (87.5% of eggs) for the former and July-December (96.0% of eggs) for the latter. The highest peaks of laying were in October and August, respectively. Kiwi eggs were considered to belong to the same clutch if they were laid no more than 40 days apart. Clutch size in the North Island brown and little spotted kiwi was most commonly one or two eggs. In the great spotted kiwi only clutches of one egg were found. The mean clutch size of North Island brown kiwi outdoors (1.51±0.05) did not differ from that of free-living North Island brown kiwi (1.33±0.09). Similarly, there was no difference in the mean clutch size of captive and free-living little spotted kiwi (1.39±0.14 and 1.10±0.07). The mean interval between eggs for kiwi outdoors did not differ between North Island brown kiwi and little spotted kiwi (27.4±0.5 and 30.7±1.8 days). The results indicate that all three species of kiwi maintain annual cycles of egg-laying in captivity, with the main egg-laying season being longer than in free-living birds.     

Ancient DNA Analyses Reveal Contrasting Phylogeographic Patterns amongst Kiwi (Apteryx spp.) and a Recently Extinct Lineage of Spotted Kiwi

The little spotted kiwi (Apteryx owenii) is a flightless ratite formerly found throughout New Zealand but now greatly reduced in distribution. Previous phylogeographic studies of the related brown kiwi (A. mantelli, A. rowi and A. australis), with which little spotted kiwi was once sympatric, revealed extremely high levels of genetic structuring, with mitochondrial DNA haplotypes often restricted to populations. We surveyed genetic variation throughout the present and pre-human range of little spotted kiwi by obtaining mitochondrial DNA sequences from contemporary and ancient samples. Little spotted kiwi and great spotted kiwi (A. haastii) formed a monophyletic clade sister to brown kiwi. Ancient samples of little spotted kiwi from the northern North Island, where it is now extinct, formed a lineage that was distinct from remaining little spotted kiwi and great spotted kiwi lineages, potentially indicating unrecognized taxonomic diversity. Overall, little spotted kiwi exhibited much lower levels of genetic diversity and structuring than brown kiwi, particularly through the South Island. Our results also indicate that little spotted kiwi (or at least hybrids involving this species) survived on the South Island mainland until more recently than previously thought.     

Serum prolactin and testosterone levels in captive and wild brown kiwi (Apteryx mantelli) during the prebreeding,breeding, and incubation periods

In brown kiwi (Apteryx mantelli), the male is the primary incubator, a trait that is relatively rare among birds. The maintenance of avian incubation behavior is controlled by the protein hormone prolactin (PRL). Although steroid hormone concentrations in both wild and captive kiwi have previously been reported, this study is the first to report levels of PRL in captive and wild male and female kiwi through the prebreeding and breeding seasons, and to directly compare testosterone (T) concentrations between captive and wild males during the breeding and incubation periods. Female PRL concentrations increased at the time of oviposition, whereas male PRL concentrations rose gradually between the prebreeding and incubation periods. Although males are considered the main incubator, an increase in PRL levels could help females maintain behaviors such as nest guarding, or to take over incubation the event of mate loss. A gradual increase in PRL allows the male to be ready for incubation during the long breeding season. Interestingly, T concentrations in captive males did not decrease during incubation and was significantly higher than in wild males. Continual elevated T could have an impact on sperm production through negative feedback, thereby contributing to the low egg fertility seen in captive kiwi. Therefore, determining the underlying reason for the differences in hormone levels could be significant, if not vital, for improving the success of captive kiwi breeding programs.     

Characterisation of class II B MHC genes from a ratite bird,the little spotted kiwi (<Emphasis Type="Italic">Apteryx owenii</Emphasis>)

Major histocompatibility complex (MHC) genes are important for vertebrate immune response and typically display high levels of diversity due to balancing selection from exposure to diverse pathogens. An understanding of the structure of the MHC region and diversity among functional MHC genes is critical to understanding the evolution of the MHC and species resilience to disease exposure. In this study, we characterise the structure and diversity of class II MHC genes in little spotted kiwi Apteryx owenii, a ratite bird representing the basal avian lineage (paleognaths). Results indicate that little spotted kiwi have a more complex MHC structure than that of other non-passerine birds, with at least five class II MHC genes, three of which are expressed and likely to be functional. Levels of MHC variation among little spotted kiwi are extremely low, with 13 birds assayed having nearly identical MHC genotypes (only two genotypes containing four alleles, three of which are fixed). These results suggest that recent genetic drift due to a species-wide bottleneck of at most seven birds has overwhelmed past selection for high MHC diversity in little spotted kiwi, potentially leaving the species highly susceptible to disease.     

Egg rejection by Iberian azure-winged magpies Cyanopica cyanus in the absence of brood parasitism

The Iberian azure-winged magpie Cyanopica cyanus shows a remarkable ability to discriminate against great spotted cuckoo Clamator glandarius eggs. Here, I studied whether egg recognition in this species could be a derived feature resulting from intra-specific brood parasitism. Azure-winged magpies showed a very high level of discrimination and rejection of great spotted cuckoo models (73.7%), and of conspecific eggs (42.8%), even when no evidence of great spotted cuckoo or conspecific brood parasitism has been found in the population. Azure-winged magpie discriminated more readily than magpies, the current favourite host of the great spotted cuckoo. The high rejection rate of conspecific eggs by the azure-winged magpie suggests that it is quite possible that egg discrimination in this species evolved in response to conspecific brood parasitism rather than to cuckoo parasitism.     

Pre- and postnatal energetics of the North Island brown kiwi (Apteryx mantelli)

In four eggs and four chicks of the North Island brown kiwi (Apteryx mantelli) we measured pre- and postnatal oxygen consumption rate (VO(2)) daily from day (d)-75 (prior to hatching) until d+25 (after hatching). The increase of embryonic VO(2) reaches a plateau phase between d-22 and d-5 (0.113 ml O(2)/g/h=59.6% of allometrically expected value of a typical 416-g egg). Mean total O(2) uptake per egg (43.01 l O(2)) corresponds to an energy turnover rate of 2.04 kJ/g during embryonic development. This is nearly identical to the expected value for all birds (2.00+/-0.8 kJ/g). Hence, the kiwi neither 'gained nor lost energy' (Calder, 1979.Biosci. 8, 461-467) by its extreme prolongation of incubation time; it is as efficient as other avian embryos. The kiwi embryo expends only approximately 17% (847 kJ) of the energy originally stored in the egg (4942 kJ). Forty-eight percent of the egg's initial yolk mass can be found as spare yolk in the hatchling and can serve as the chick's sole source of energy and substrate for tissue production for up to at least 17 days after hatching.     

Assessment of reproductive status and ovulation in female brown kiwi (Apteryx mantelli) using fecal steroids and ovarian follicle size

Despite being listed as endangered on the IUCN (World Conservation Union) red list [BirdLife International 2005], and intense conservation efforts on its behalf, little is known about the reproductive physiology of the brown kiwi (Apteryx mantelli). This study therefore was undertaken to describe ovarian function in brown kiwi using noninvasive methods that can be utilized in the field. This study describes a simple method for extraction and detection of steroid metabolites in brown kiwi feces by enzyme‐linked immunosorbent assay (ELISA). Average fecal steroid hormone metabolite concentrations were derived from 10 egg‐lay cycles in 2 hens. Testosterone began to increase between 23 and 18 days before oviposition (?23d to ?18d) and remained elevated until just before oviposition. Estradiol rose from ?28d to ?13d and remained elevated until 12 days after oviposition. Progesterone exhibited an increase of short duration from ?8d to +7d with a peak at ?3d. Ovarian follicle growth rate was determined by serial ultrasound measurements of follicles in three habituated female kiwi. Follicle growth rate was described by the formula y=132.54e^0.0569x (y=follicle diameter, x=days before oviposition). Follicles ranged in size from 10 to 76 mm. Day of ovulation was estimated using egg yolk size, follicle growth curve, and day of oviposition. This yielded an ovulation‐oviposition cycle length of 7–10 days, significantly shorter than the only other estimate of 16–17 days derived using body weights of a single kiwi hen. The correlation of fecal steroid hormone concentrations with follicle growth provides new insight into the reproductive physiology of the female brown kiwi. Zoo Biol 0:1–10, 2005. © 2005 Wiley‐Liss, Inc.     

Eavesdropping cuckoos: further insights on great spotted cuckoo preference by magpie nests and egg colour

Reproductive success of brood parasites largely depends on appropriate host selection and, although the use of inadvertent social information emitted by hosts may be of selective advantage for cuckoos, this possibility has rarely been experimentally tested. Here, we manipulated nest size and clutch colouration of magpies (Pica pica), the main host of great spotted cuckoos (Clamator glandarius). These phenotypic traits may potentially reveal information about magpie territory and/or parental quality and could hence influence the cuckoo’s choice of host nests. Experimentally reduced magpie nests suffered higher predation rate, and prevalence of cuckoo parasitism was higher in magpie nests with the densest roofs, which suggests a direct advantage for great spotted cuckoos choosing this type of magpie nest. Colouration of magpie clutches was manipulated by adding one artificial egg (blue or cream colouration) at the beginning of the egg-laying period. We found that host nests holding an experimental cream egg experienced a higher prevalence of cuckoo parasitism than those holding a blue-coloured egg. Results from these two experiments suggest that great spotted cuckoos cue on magpie nest characteristics and the appearance of eggs to decide parasitism, and confirm, for the first time, the ability of cuckoos to distinguish between eggs of different colours within the nest of their hosts. Several hypothetical scenarios explaining these results are discussed.     

Ancient DNA and conservation: lessons from the endangered kiwi of New Zealand

Conservation genetics typically seeks to map the distribution of contemporary genetic variation across space, and to use the resulting genetic parameters to infer any likely short-term evolutionary consequences for rare and endangered species. Recent developments in the study of ancient DNA now enable the extension of genetic variation studies backwards in time and provide a context by which to interpret contemporary levels of genetic variation, in addition to any patterns of genetic change over time. Ancient DNA research can also help to determine past levels of genetic diversity, identify species' boundaries and reveal former ranges among morphologically cryptic taxa. Ancient DNA sequence data for the New Zealand kiwi (Apteryx spp.) are presented and we show that most ancient populations of rowi and tokoeka exhibited private mitochondrial DNA haplotypes. Moreover, we illustrate that the extinction of these populations has led to the loss of considerable genetic variation. We also use ancient DNA methodology to determine past species distributions for brown kiwi and great spotted kiwi whose bones are morphologically indistinguishable.     

The costs of egg production and incubation in great tits (Parus major).

The costs of egg production and incubation may have a crucial effect on avian reproductive decisions, such as clutch size and the timing of reproduction. We carried out a brood-size enlargement experiment on the great tit (Parus major), in which the birds had to lay and incubate extra eggs (full costs), only incubate extra eggs (free eggs) or did not pay any extra cost (free chicks) in obtaining a larger brood. We used female fitness (half the recruits produced plus female survival) as a fitness measure because it is the female which pays the costs of egg production and incubation, and because clutch size is under female control. Female fitness decreased with increasing costs (fitness of free chicks females is higher than that of free eggs females which is higher than that of full costs females). These fitness differences were due to differences in female survival rather than in the number of recruits produced. This is the first time that the costs of egg production and incubation have been estimated using such a complete fitness measure, including, as our measure does, the local survival to the following year of both the female and her offspring. Our results emphasize that reproductive decisions cannot be understood without taking egg production and incubation costs into account.     

Kiwialges haastii n. sp., a feather mite (Acari: Analgidae) from the great spotted kiwi,Apteryx haastii Potts, 1872 (Aves: Apterygidae); with a key to species of Kiwialges,and new host records

Abstract

A new species of feather mite (Acari: Analgidae), Kiwialges haastii n. sp. is described from the great spotted kiwi, Apteryx haastii Potts, 1872. Four new host records are reported and a key to species of Kiwialges is presented.     

Breeding biology aspects of spotted flapshell Turtle,Lissemys punctata (Lacepede 1788), in Bangladesh

This study was conducted to determine the breeding season, gonadal development, egg laying period, clutch size and other biological aspects of spotted flapshell turtle, Lissemys punctata, of Bangladesh between January 1997 and December 2001. The egg laying period of L. punctata, was found between August and March. The nesting sites were elevated fallow lands in secluded areas. The female turtle laid all the mature eggs at a time for each clutch at night. A gravid female turtle laid three clutches of eggs in each year and the mean clutch size was 13.0 ± 1.9 eggs and mean weight of each egg was 10.3 ± 1.3 g. The eggs are spherical in shape and whitish in color. The mean incubation period was 173 ± 34 days (range 119–225 days). The incubation period of first clutch was the longest than the second and third clutches. Hatching success was found 41%. Maximum hatching was observed in June. The present investigation was made to explore the possibility to raise turtle farming in captive condition. The findings would, hopefully, help to rear the species and to assess the commercial potentiality of turtle farming in captive condition, that is, in the eco-climatic condition of Bangladesh.     

A new method for catching cavity‐nesting birds during egg laying and incubation

ABSTRACT The physiological condition of female birds during the egg‐laying and incubation periods is of considerable interest and yet is relatively understudied in wild birds, primarily due to the difficulty of catching birds during this period without causing nest desertion. We therefore developed a box‐net to capture cavity‐nesting birds using sections of a mist‐net placed around a metal cubic frame. We captured female Great Tits (Parus major) as they left nest boxes during the egg‐laying and incubation periods and measured desertion rates. Using box‐nets, we captured 108 of 119 (90%) females during egg laying and 10 of 12 (83%) during incubation. Our recapture rate over two consecutive days during incubation was 50% (5 of 10). Females not captured left nest boxes before we attempted to capture them, escaped through a hole in the mist‐net, or remained in nest boxes for more than 2 h, after which we ended capture attempts. Overall, 22% of egg‐laying females deserted, with desertion rates highest early in the egg‐laying period. Desertion rates of females captured using box‐nets did not differ from those of undisturbed females. One of 10 females captured in a box‐net deserted during the incubation period. Box‐nets are portable, can be set up and taken down quickly and easily, and could potentially be used with nest boxes or natural cavities at any height. Box‐nets are easy to construct and adaptable for use with an array of cavity‐nesting birds, and can be an important tool for studying female physiology during egg laying and incubation.     

自然条件下福寿螺繁殖特性

为了弄清福寿螺在自然条件下的繁殖特性,在湖南省资兴市水稻田内进行了为期3年的人工养殖试验.结果表明:福寿螺在湘南自然条件下1年可发生不完全3代.第1代、第2代和第3代(越冬代)雌螺的平均性成熟期分别为59.3、45.4和213.0 d.雄螺的平均性成熟期比雌螺早4.3d.福寿螺的自然性比(♀∶♂)为1.54:1.每次交配持续时间平均为19.2h,但并非每次交配后都会产卵.田间福寿螺月产卵块数与月平均气温呈显著正相关(r=0.756).福寿螺卵块的平均孵化期(d)与日平均气温(℃)呈显著负相关(r=-0.980),其线性回归方程为y=-0.726x+23.064.福寿螺卵块的平均孵化期为20.7d,平均孵化率为44.1％.雌螺与雄螺的平均寿命分别为2.40和1.98 a.平均每头雌螺一生可产卵13764粒,可繁殖幼螺6070头.     

Rejection of parasitic eggs in relation to egg appearance in magpies

The coevolutionary process between avian brood parasites and their hosts predicts that low intraclutch variation in egg colour appearance favours egg discrimination of parasite eggs by hosts. Low intraclutch variation would also result in high interclutch variation, which would increase the difficulty of evolution of mimicry by the cuckoo, because many host colour patterns might coexist in the same host population. We explored this possibility using an experimental approach in the common magpie, Pica pica, and great spotted cuckoo, Clamator glandarius, system. We artificially parasitized magpie nests with great spotted cuckoo model eggs to assess host response in two populations in Spain (Guadix and Doñana) in relation to intraclutch variation in egg appearance, measured by ultraviolet-visible reflectance spectrophotometry. Individuals that rejected model cuckoo eggs had higher intraclutch variation than accepters, suggesting that an increase, rather than a decrease, in intraclutch variation in magpie egg appearance was advantageous for cuckoo egg discrimination.     

Eggshell Spottiness Reflects Maternally Transferred Antibodies in Blue Tits

Blue-green and brown-spotted eggshells in birds have been proposed as sexual signals of female physiological condition and egg quality, reflecting maternal investment in the egg. Testing this hypothesis requires linking eggshell coloration to egg content, which is lacking for brown protoporphyrin-based pigmentation. As protoporphyrins can induce oxidative stress, and a large amount in eggshells should indicate either high female and egg quality if it reflects the female''s high oxidative tolerance, or conversely poor quality if it reflects female physiological stress. Different studies supported either predictions but are difficult to compare given the methodological differences in eggshell-spottiness measurements. Using the blue tit Cyanistes caeruleus as a model species, we aimed at disentangling both predictions in testing if brown-spotted eggshell could reflect the quality of maternal investment in antibodies and carotenoids in the egg, and at improving between-study comparisons in correlating several common measurements of eggshell coloration (spectral and digital measures, spotted surface, pigmentation indices). We found that these color variables were weakly correlated highlighting the need for comparable quantitative measurements between studies and for multivariate regressions incorporating several eggshell-color characteristics. When evaluating the potential signaling function of brown-spotted eggshells, we thus searched for the brown eggshell-color variables that best predicted the maternal transfer of antibodies and carotenoids to egg yolks. We also tested the effects of several parental traits and breeding parameters potentially affecting this transfer. While eggshell coloration did not relate to yolk carotenoids, the eggs with larger and less evenly-distributed spots had higher antibody concentrations, suggesting that both the quantity and distribution of brown pigments reflected the transfer of maternal immune compounds in egg yolks. As yolk antibody concentrations were also positively related to key proxies of maternal quality (egg volume, number, yellow feather brightness, tarsus length), eggshells with larger spots concentrated at their broad pole may indicate higher-quality eggs.     

Molecular and other biological evidence supports the recognition of at least three species of brown kiwi

The presence of morphologically crypticlineages within the threatened brown kiwi ofNew Zealand has confounded their taxonomy. Recent genetic studies (Herbert and Daugherty1994; Baker et al. 1995) revealed that atleast two phylogenetic species exist within thebrown kiwi, and suggested that further researchshould resolve the taxonomic problems. In thispaper we extend genetic analyses to includesequences from 58 brown kiwi representing fivephylogenetic lineages for four mitochondrialloci (control region, cytochrome b,ATPase 6 and ATPase 8). Major lineages ofbrown kiwi are shown to be reciprocallymonophyletic, and align with other biologicaldifferences in the ecology, behavior,morphology and parasites of kiwi. BecausemtDNA sequences of major lineages of kiwi arenot evolving in a clocklike manner, we used anew penalized likelihood method withrate-smoothing to date the divergence of NorthIsland brown kiwi and the geographicallyisolated Okarito population (rowi) at about 6.2Mya. These lineages diverged about 8.2 Myafrom the brown kiwi in Fiordland and Haast inthe southern part of the South Island, and arethus older than the species of spotted kiwi(5.8 Mya). Given their distinctness, long-termgeographical isolation, lack of hybridizationin introduced populations, and accumulation ofnew biological characters within theselineages, we hypothesize that reproductiveincompatibilities have probably arisen as well. We therefore recommend that these divergentlineages be formally recognized as fullspecies; Apteryx mantelli should bere-instated for the North Island brown kiwi,A. australis should be restricted to thetokoeka, and a new species A. rowiishould be erected to describe the rowi atOkarito. Tokoeka should be split into at leastthree conservation management units (Haast,Fiordland and Stewart Island [Rakiura]), butfurther research is required to determine theexact relationships and status of theselineages. Further investigation is alsorequired into the genetic structuring of theNorth Island brown kiwi to confirm conservationmanagement units on the North Island.     

A male destroying an egg in a cooperative breeding attempt in Lesser Spotted Woodpecker Dendrocopos minor

A cooperative breeding case was found in Lesser Spotted Woodpecker (LSW) in which a male and three different females were involved. The contributions of each member of the cooperative breeding team were quantified during nest excavation and incubation. This is the first such report on LSW. During daytime, the nest was mainly occupied by the male during the final excavation phase, nest guarding, and egg laying. Aggressive interactions were recorded between two of the females during nest excavation, egg laying and incubation. The male was recorded destroying an egg presumably laid by the dominant female. The highest contribution during daytime incubation was made by the dominant female (39%), followed by the male (34%) and then by a second female (27%). The third female contributed very little at any stage. A picture of the male removing the egg was analyzed to estimate the egg maximum width, ruling out a possible runt egg. Motivations behind the behaviour of the male destroying the egg remain unclear.