The behavioral endocrinology of domestication: A comparison between the domestic guinea pig (Cavia aperea f. porcellus) and its wild ancestor, the cavy (Cavia aperea)

In this study spontaneous behavior and endocrine parameters were compared between the domestic guinea pig (Cavia aperea f. porcellus) and its wild ancestor, the cavy (Cavia aperea), to elucidate the process of domestication in this species. In 120 h of observation time the behavior of five groups of wild and seven groups of domestic guinea pigs, each consisting of one adult male and two adult females, was analyzed quantitatively. To assess the activities of the pituitary-adrenocortical (PAC), the pituitary-gonadal (PG), and the sympathetic-adrenomedullary (SAM) systems, serum cortisol, testosterone, epinephrine, and norepinephrine concentrations, as well as adrenal tyrosine hydroxylase activities, were determined in males of both forms. The following significant differences between wild cavies and domestic guinea pigs were found: the domesticated animals displayed less aggressive but more sociopositive and more male courtship behavior than their wild ancestors. In addition, they were distinctly less attentive to their physical environment than the wild cavies. The basal activity of the SAM system, as well as the reactivity of the SAM and the PAC systems, was distinctly reduced in the domesticated animals. In contrast, the basal activity of the PAC system did not differ between both forms. The activity of the PG system was significantly higher in males of the domestic guinea pig than in male wild cavies. Thus, in guinea pigs the process of domestication has led to typical behavioral traits-reduced aggressiveness, increased social tolerance-which have also been found in comparisons between wild and domestic forms of other species. The decreased reactivity of the organism's stress axes can be regarded as a physiological mechanism which helps domesticated animals to adjust to man-made housing conditions.     

Body Weight and Rearing Conditions of Males, Female Choice and Paternities in a Small Mammal, Cavia aperea

Patterns of mate choice may be important determinants of a species' social organisation and mating system. At least two different aspects of female mate choice can be distinguished: choice of a social partner and choice of the genetic father of the offspring. Different characteristics of males can qualify them for these two roles. Although social and reproductive partners have been shown to differ in many species, social association times are often used in laboratory choice tests to infer reproductive preferences. The traits for which females may choose partners are diverse. Body size can correlate with the male's strength in defending resources or other abilities benefiting the female and her offspring. In species living in social groups, social skills learned from group members during infancy can be important for later reproductive success. In this laboratory study, we conducted choice tests with wild cavies, Cavia aperea , a harem-living species of South American rodents, to determine social preferences of females towards two simultaneously available males. For offspring sired during these tests, paternities were determined by microsatellite DNA profiling. Males used in the tests differed in body weight and in rearing conditions: Half of the males had been reared in the presence or absence of their father, respectively. Male rearing conditions had no effect on either female social preferences or paternities. Females significantly preferred heavier males as social partners. In five of six tests, the heavier male also sired the offspring. Sires were in most cases but not consistently socially preferred. Heavier males may be preferable as social partners because they are better able to provide females with resources or have more experience in paternal care or predator avoidance as weight correlates with age. When choosing reproductive partners, females may prefer other male traits and the distribution of paternities may also be influenced by sperm competition.     

Photoperiod influences the development and the expression of personality traits and social behaviour in wild cavies (Cavia aperea)

Seasonality influences behaviour, physiology and life history of organisms, and photoperiod is a reliable and influential cue gearing individual adaptation to seasonal changes in the environment. Plastic phenotypic adjustments according to the prevailing season are a widespread, well-studied phenomenon. Less well known are effects on developmental trajectories of animals born into different seasons. By manipulating the photoperiod during pregnancy and lactation independent of food availability and temperature, we studied if maternal behaviour in the precocial wild cavy (Cavia aperea) responds to this seasonal cue and if maternal behaviour and photoperiod shape offspring development and behaviour. We repeatedly observed mothers before and after birth. We investigated if pups differ in risk-taking behaviours directly after birth (indicating direct prenatal effects) or only after gaining independence of the mother (indicating delayed prenatal or postnatal effects due to own cue sampling). Furthermore, we tested if different simulated seasons of birth affect social and aggressive behaviour of adult animals. While theoretical predictions according to risk-taking are quite clear, they are contradictory for social behaviours that are much less well studied. We predicted that animals born into spring photoperiods would be aggressive because of a need to gather enough resources for early reproduction and successfully establishing a territory at least in males. Spring-born males were more aggressive than autumn-borns, while females did not differ with respect to season. Spring-born females, however, lost more body mass when they had to integrate into an established group and autumn-born females initiated more aggressive interactions. Our data demonstrate flexible and long-term stable seasonal effects on behaviours that likely affect fitness outcomes under natural conditions.     

Maternal Effort is State Dependent: Energetic Limitation or Regulation?

Many small altricial rodents have a postpartum oestrus and are often simultaneously pregnant and lactating. Negative influences of concurrent pregnancy and lactation on both lactational performance and the litter in utero are commonly observed and have been interpreted as resulting from high simultaneous energetic demands of gestation and lactation. We studied these effects in the precocial guinea‐pig (Cavia aperea f. porcellus) that, like many altricial rodents, has a postpartum oestrus, but in which the peaks of energy expenditure on lactation and gestation are widely separated. This life history allowed to investigate whether physiological regulation other than by energetic limitations may be responsible for allocation conflicts arising when lactation and gestation overlap. By comparing simultaneously pregnant and lactating females with lactating non‐pregnant females, we show that females in the former group nurse less and wean earlier than females of the latter group. In a comparison of litter size, litter mass, and pup mortality of females that had not been lactating during pregnancy with females that had been simultaneously pregnant and lactating, we show that the latter do not reduce investment in the following litter. In our study, energetic constraints on ad libitum fed females are unlikely and we therefore suggest that the results must be explained by regulatory constraints on lactational effort. We point out that this explanation has not been excluded for the effects observed in altricial small mammals.     

Is a wild mammal kept and reared in captivity still a wild animal?

This study compared domestic guinea pigs (Cavia aperea f. porcellus; DGP) and two different populations of the wild cavy (Cavia aperea), its ancestor, to examine whether rearing of wild mammals in captivity affects their behavior and physiological stress responses. One population of wild cavies consisted of wild-trapped animals and their first laboratory-reared offspring (WGP-1). The animals of the other population were reared in captivity for about 30 generations (WGP-30). The spontaneous behavior of each of six groups of WGP-1 and WGP-30 and nine groups of DGP, each consisting of one adult male and two adult females, was analyzed quantitatively. Blood samples of the males were taken to determine cortisol, epinephrine, and norepinephrine concentrations. In addition, the exploratory behavior of 60-day-old male WGP-1, WGP-30, and DGP was investigated in an exploration apparatus. The domesticated animals displayed significantly less aggression, but significantly more sociopositive and male courtship behavior than their wild ancestors. In addition, DGP were much less attentive to their physical environment. Surprisingly, no behavioral difference was found between WGP-1 and WGP-30. Basal cortisol concentrations did not differ between wild and domestic guinea pigs. Catecholamine concentrations, however, as well as the challenge values of cortisol, were distinctly reduced in the DGP. WGP-1 and WGP-30 did not differ with respect to their endocrine stress responses. In the exploration apparatus both forms of wild cavies were much more explorative than the domestic animals. These data suggest that the long-term breeding and rearing of wild guinea pigs in captivity do not result in significant changes in behavior and hormonal stress responses. It appears to take much longer periods of time and artificial selection by humans to bring about characters of domestication in wild animals.     

Senescence and costs of reproduction in the life history of a small precocial species

Species following a fast life history are expected to express fitness costs mainly as increased mortality, while slow‐lived species should suffer fertility costs. Because observational studies have limited power to disentangle intrinsic and extrinsic factors influencing senescence, we manipulated reproductive effort experimentally in the cavy (Cavia aperea) which produces extremely precocial young. We created two experimental groups: One was allowed continuous reproduction (CR) and the other intermittent reproduction (IR) by removing males at regular intervals. We predicted that the CR females should senesce (and die) earlier and produce either fewer and/or smaller, slower growing offspring per litter than those of the IR group. CR females had 16% more litters during three years than IR females. CR females increased mass and body condition more steeply and both remained higher until the experiment ended. Female survival showed no group difference. Reproductive senescence in litter size, litter mass, and reproductive effort (litter mass/maternal mass) began after about 600 days and was slightly stronger in CR than IR females. Litter size, litter mass, and offspring survival declined with maternal age and were influenced by seasonality. IR females decreased reproductive effort less during cold seasons and only at higher age than CR females. Nevertheless, offspring winter mortality was higher in IR females. Our results show small costs of reproduction despite high reproductive effort, suggesting that under ad libitum food conditions costs depend largely on internal regulation of allocation decisions.     

New findings on the origin of Cavia intermedia,one of the world's rarest mammals

Cavia intermedia, a rare species of cavy found exclusively on Moleques do Sul Island, Brazil, differs from other cavies in genetic and morphological characteristics and has been regarded as phylogenetically closest to Cavia magna. Here, I present morphological evidence for the likely origin of Cavia intermedia from an ancestor shared with Cavia magna: an external female genital organ similar in size and appearance to the male's penis. Masculinization of the female genitalia has been observed in Cavia magna but does not occur in Cavia aperea, which suggests a sister relationship between Cavia magna and Cavia intermedia.     

Olfactory Individual Recognition of Mothers by Young Guinea-Pigs (Cavia porcellus)

In mammals, olfactory cues play a major role in individual recognition and urine is one source of potentially individual‐specific olfactory cues. We studied how soon young of the extremely precocial domestic guinea‐pig (Cavia porcellus) establish specific preferences for maternal urine smell by offering 5–30‐d‐old young a simultaneous choice between a urine sample of the mother and urine from an unfamiliar unrelated lactating female. Young showed increasing preference for the smell of maternal urine from day 5 of life onwards. On day 10 of life, they discriminated between maternal urine and that of other lactating females when these were unfamiliar and related, unfamiliar and unrelated or familiar unrelated, but not when the urine of the other female came from a familiar and related lactating animal. The last result is based on fewer litters and, therefore has to be considered as preliminary. As our results are based on spontaneous preferences for just one source of olfactory cues, discrimination of live animals is likely to be even better than demonstrated here. Learning or phenotype matching of individual specific cues enable these precocial young to form a specific bond with their mother soon after birth.     

Timing of reproduction in a Darwin''s finch: temporal opportunism under spatial constraints

We investigated whether predation by the minor grison ( Galictis cuja , a small mustelid) played a key role in limiting a wild cavy population ( Cavia magna ), ultimately leading to its local extinction. Radio-telemetry and capture-mark-recapture techniques were used to estimate grison predation rates (kill rates), time-specific probabilities of apparent mortality (population loss rate), overall mortality and grison predation for the cavy population. Additionally, we present data on alternative prey species, grison diet and reproduction to show potential proximate mechanisms of grison predation on wild cavies. The predictions specified were mostly confirmed: (1) grison predation was responsible for almost 80% of the cavies killed by known predators; (2) grison predation probabilities paralleled those of overall mortality of cavies over time; and (3) also those of the apparent mortality of the population. Thus, the population dynamics and the local extinction of the cavy population were not due to emigration processes. (4) Grison predation rates were not density-dependent, but showed pronounced peaks during the austral summer. The grison mainly preyed on small mammals: two water-rat species and the wild cavies. When the availability of alternative prey decreased in summer, the grison appeared to specialise on cavies. The onset of grison reproduction was somewhat delayed in relation to the onset of cavy reproduction. The lack of alternative prey coincided with high grison food demands due to reproduction, leading to a very high predation pressure ultimately resulting in the local extinction of the cavy population. We conclude that grison predation was indeed the main factor driving changes of the cavy population studied and speculate why caviomorph rodents might be especially susceptible to local extinction processes.     

阿根廷淡水大米草沼泽中野生豚鼠的种群统计和微生境的利用

本文研究了秋冬季节淡水大米草(Spartina densiflora)沼泽中野生豚鼠(Cavia a perea)的丰度、繁殖和微生境利用,以及其对当地植被和棉鼠类(Sigmodontine)啮齿动物的影响。野生豚鼠喜好S.densiflora覆盖度高的生境。繁殖个体(成体)主要利用矮草为主的斑块,幼体则主要利用禾本科植物为主的斑块。结果说明,野生豚鼠微生境的利用受捕食风险和食物种类的影响。在淡水沼泽中,野生豚鼠的丰度、繁殖、体重和微生境利用没有季节性变化,它们对植物的取食和活动跑道的建造对植被结构和同域的啮齿类动物没有负面影响。通过对具有中度季节性变化的淡水生境中的豚鼠种群和具有高度季节变化的草地和路边中的豚鼠种群进行比较,表明野生豚鼠的种群动态以及豚鼠种群对植被和与其共生的啮齿动物群落的影响都受到冬季植被盖度的限制。     

Does the unusual life history of the precocial cavy (Cavia magna) translate into an exceptional field metabolic rate?

An association between metabolic rate and life history has been predicted but not demonstrated yet. To test this prediction, we measured the field metabolic rate (FMR) of a rodent, Cavia magna, which has an unusual combination of life-history traits. Cavies live aboveground, feed on energy-poor diets, and take relatively long to produce a few highly precocial young. This "slow-living" life history might predict a relatively low metabolic rate. We recorded FMR of 29 individuals of C. magna in Uruguay using doubly labeled water. This represents the first FMR record for a hystricomorph rodent. Besides body mass, season best explained variation in recorded FMR, with higher daily energy expenditures during winter. Surprisingly, the reproductive state of females did not affect FMR. Comparison with allometric prediction did not support the hypothesis that the slow-living life history of precocial grazing cavies translates into relatively low FMR. However, the clear effect of season on FMR as well as the fact that cavies appear able to compensate for the energy cost of reproduction might be associated with the nonburrowing precocial life history of this species.     

Reproductive biology,growth and age composition of the introducedOreochromis mossambicus (Cichlidae) in two reservoirs,Brisbane, Australia

Synopsis The reproductive biology, growth and age composition ofOreochromis mossambicus (Peters) were studied in two Brisbane reservoirs during 1981–1983. The life history was assessed in terms of altricial and precocial tendencies after Noakes & Balon (1982). Fish from both reservoirs had a precocial life history style, showing delayed maturation at 12–15 months, at minimum sizes of 152 mm standard length in males and 174 mm standard length in females. The minimum size of fish at maturity was 0.51–0.73 of maximum size. Mean fecundity was 1360 (± 117.2 standard error) in North Pine Dam and 2107 (220.4 standard error) in Tingalpa Reservoir. This difference was significant (p<0.01). Oöcyte diameter ranged from 1.7–2.6 mm. Overall sex ratio was 1:1 but males predominated amongst fish larger than 200 mm standard length. Stocks consisted mainly of three age groups and the oldest fish caught were 3+ years old. Growth rates were high in both populations. Length-weight relationships conformed to the allometric formula W = aLⁿ and were consistent with data on growth rates. High growth rates are attributed to the high productivity of both reservoirs and the small size of the populations relative to food resources, which are not utilized directly by other fishes. These factors probably also account for the precocial life history style exhibited, compared to the altricial life history style ofO. mossambicus in Lake Sibaya, South Africa, a food-restricted environment.     

Reproduction in captive female Cape porcupines (Hystrix africaeaustralis)

Captive females attained sexual maturity at an age of 9-16 months and conceived for the first time when 10-25 months old. Adult females were polyoestrous but did not cycle while lactating or when isolated from males. The length of the cycle varied from 17 to 42 days (mean +/- s.d. 31.2 +/- 6.5 days; n = 43) and females experienced 3-7 sterile cycles before conceiving. Pregnancy lasted for 93-94 days (93.5 +/- 0.6 days; N = 4) and litter intervals varied from 296 to 500 days (385 +/- 60.4; n = 10). Litter size varied from 1 to 3 (1.5 +/- 0.66; n = 165) and the well-developed precocial young weighed 300-440 g (351 +/- 47.4 g; n = 19) at birth. Captive females reproduced throughout the year with most litters (78.7%; n = 165) being produced between August and March.     

Reproductive biology and postnatal development in the tent-making bat Artibeus watsoni (Chiroptera: Phyllostomidae)

In this study we investigated the reproductive patterns and postnatal development in the tent-making bat Artibeus watsoni . We sampled two populations in the Golfito Wildlife Refuge and Corcovado National Park, south-western Costa Rica, from June 2003 to March 2005. Most females were pregnant during the months of January and June, and most were lactating in March and July, indicating that this species exhibits seasonal bimodal polyoestry, with the first parturition peak occurring in February–March and the second in June–July. Additionally, we observed a postpartum oestrus following the first parturition, but not after the second. Females entered oestrus again in November–December and had a gestation period of c . 3 months. A female-biased sex ratio of neonates was observed during the second parturition period, and young were born at 32 and 56% of their mothers' body mass and length of forearm, respectively. Adult proportions in length of forearm were attained faster than adult proportions in body mass, and sustained flight was only possible after 35 days of age, when pups had achieved 100 and 80% of adult length of forearm and body mass proportions, respectively. Weaning and roosting independence occurred when young were c . 30–40 days old, and young females appeared to remain close to their place of birth, at least for their first mating period, whereas adult males were never recaptured near their birth site. In addition, sexual maturity was reached in as little as 3 months in females born during the first parturition period, whereas females born during the second birth period in June–July seemed to reach maturity at 6 months of age. Our results show that A. watsoni belongs to the faster lane of the slow–fast continuum of life-history variation in bats, which may be attributed primarily to its roosting and feeding ecology.     

Animal Housing and Welfare: Effects of Housing Conditions on Body Weight and Cortisol in a Medium-Sized Rodent (Cavia aperea)

Rodents are the most abundant experimental nonhuman animals and are commonly studied under standard laboratory housing conditions. As housing conditions affect animals' physiology and behavior, this study investigated the effects of indoor and outdoor housing conditions on body weight and cortisol level of wild cavies, Cavia aperea. The changing housing condition strongly influenced both parameters, which are commonly used as indicators for animal welfare. The transfer from outdoor to indoor enclosures resulted in a body-weight loss of about 8%. In contrast, animals kept indoors showed a substantial weight gain of about 12% when they were transferred outdoors. These effects were reversible. To substantiate a connection between body-weight changes and the health states of the animals, blood basal cortisol concentrations were measured. Animals kept outdoors had significantly lower cortisol levels than did animals kept indoors. These results imply that indoor conditions have a direct effect on the animals' states. The physiological and metabolic consequences as well as potential welfare aspects should be taken into account when planning experimental work, especially on nondomestic animals.     

Ontogenesis of the electroretinogram in a precocial mammal, the guinea pig (Cavia porcellus)

1. Ontogenesis of the electroretinogram, the mass electrical response of the retina to flash light stimuli, was studied in the guinea pig (Cavia porcellus), a precocial species with visual function at birth. 2. a-Wave components, b-wave, oscillatory potentials, slow PIII, and c-wave responses to flash stimuli developed between 55 and 64 days of gestation (full term is 68-69 days). 3. a-Waves attributable to photoreceptor functions were fully mature at 60 days. 4. ERG development lagged behind the reported critical milestones in retinal development; its prenatal onset indicates that no history of light entrainment is required for initiation of a mature ERG response.     

Ancient and modern steps during the domestication of guinea pigs (Cavia porcellus L.)

To test whether there are differences between living lineages of domestic guinea pigs Cavia porcellus , we studied 118 specimens from six breeds collected along six Andean countries as well as 15 from the wild cavy species ( Cavia tschudii ). The mean weight and body length of 15 adult wild cavies (295±31 g, 242±8.3 mm) were significantly smaller than 25 creole guinea pigs from Bolivia and Chile (639±157 g, 287±23.7 mm, respectively). Eighteen laboratory/pet guinea pigs (including the English Pirbright breed) were also smaller (900±173 g, 308±21 mm) than 25 improved ones from Peru (Tamborada breed, 1241±75.4 g, 317±12 mm) and Ecuador (Auqui breed, 1138±65.5 g, 307±8 mm). Similar size increases appeared in the first axis of a principal component analysis of six skeletal measurements, recovering 84% of total variation. Phylogenetic and haplotype analyses of complete cytochrome b gene sequences consistently joined all 22 domestic individuals (13 shared unambiguous substitutions, 100% bootstrap in 1000 replicates), probably from a single first ancient domestication in the western Andes. Six laboratory/pet sequences were also joined within a common branch (six shared substitutions, 96% bootstrap), probably from a documented European second phase. By contrast, those from improved Auqui joined a northern creole subgroup (one shared substitution, 84% bootstrap), and those from Nativa and improved Tamborada clustered together and with a southern creole subgroup (four shared substitutions, 86% bootstrap); this suggests at least two independent modern events during a more complex third phase, producing two improved guinea pigs selected for size and meat. Cavia tschudii sequences showed some unexpected geographic variation.     

Evidence from the oestrous cycle for male-induced ovulation in Bettongia penicillata (Marsupialia).

Female brush-tailed bettongs (Bettongia penicillata) housed in a breeding group of one male and one to three females had an average gestation period of 21.2 days (n = 58) and parturition was followed within 24 h by oestrus and mating. If a young was not born as a result of the mating, oestrus recurred after about 21.7 days (n = 12). From removal of pouch young to birth was 17.5 days, on average (n = 85), in females that had mated post partum, but most females that were isolated from a male before parturition returned to oestrus about 6.6 days after simultaneous removal of pouch young and return to the male (n = 9). Females housed in female-only groups appeared not to come into oestrus or to do so irregularly. These females, when returned to the male, usually came into oestrus within 10 days. These data provide evidence that, in most females of B. penicillata, ovulation does not occur in the absence of a male and that previously isolated females return to oestrus within 10 days of return to the male.     

Effect of weaning time on the growth rate and food intake of the spiny mouse pup

The spiny mouse Acomys cahirinus produces well-developed pups although it is a relatively small mammal (45 g). We envisioned two opposing hypotheses on the effect of early weaning on the growth rate of pups. The first predicts little effect since the increase in energy intake of dams above non-reproducing females is relatively low, suggesting that pups consume a large portion of their energy as solid food, and the pups are very well developed at birth. The second predicts a substantial effect since the 'index of precociality', that is the energy intake for maintenance of a pup as a proportion of that predicted for a rodent of its body mass, falls within values for altricial rodents, suggesting an extended maternal dependence of the young. To test these hypotheses, we measured the growth rate and food intake of pups weaned after either 7, 14, 21 or 28 days. Only three of 12 pups weaned after 7 days survived and, consequently, the latter hypothesis was supported. All pups weaned at 14–28 days survived. There was a significant decrease in growth rate during the first day after weaning in pups weaned at 7, 14 and 21 days but not after 28 days, suggesting that pups did not require parental care by day 28. Peak growth rate in pups weaned at 14 days occurred in the second week but occurred in the third week in pups weaned at 21 and 28 days. In spite of these differences, pups in all treatments had similar body mass at 64 days, indicating compensatory growth. We concluded that pups of A. cahirinus are precocial from a morphological aspect in that they are well developed at birth but altricial from a nutritional aspect in that they require extended maternal support.     

Morphogenesis of the cranium of Cavia porcellus L. II: Comparative part and literature

The development of the chondrocranium of Cavia porcellus is compared to those of other rodents. The tectum posterius of the investigated rodents is orientated vertically. This position is functionally caused by the attachment of the muscles of the neck and shoulder girdle. The paracondylar process is a typical feature of rodents although absent in Mesocricetus. Only in Cavia and Tatera, the connection between the lamina supraoccipitalis and the auditory capsule-the supraoccipitocapsular commissure-is missing. Youssef's (1966) generalization that the course of the notochord in rodents is of transbasal type cannot be confirmed. In Cavia, the auditory capsule is connected with the occipital region only by the exoccipitocapsular commissure. The connection between auditory capsule and basal plate is established by the alicochlear and the anterior basicapsular commissures. In comparison to other rodents, the number of commissures in Cavia is reduced. In rodents, there is always a subarcuate fossa which in later stages of development is filled out by the flocculus cerebelli. In contrary to Rajtova's (1972a) statement, Cavia shows a suprafacial commissure as all mammals do (Reinbach 1952). As the tegmen tympani is absent in Otomys and Erethizon, it is not a typical rodent feature. The carotid foramen is well developed in Cavia but the internal carotid artery obliterates until the 25 mm CRL-stage. In embryonic rodents, the ala temporalis may have a foramen ovale but not a foramen rotundum. During ontogeny rodents show the ala hypochiasmatica for the attachment of the straight muscles of the eyeball. In Cavia the ala hypochiasmatica develops independently and fuses with the postoptic root of the ala orbitalis in later stages. In myomorphs and sciumorphs, the orbitoparietal and orbitonasal commissures are present. Only in caviomorphs this part of the primary sidewall of the skull is uncomplete. Erethizon, however, shows an orbitonasal commissure whereas in Cavia both commissures are missing. In this respect the guinea-pig resembles the condition of primates. There is no interorbital septum in rodents. The nasal capsule of rodents contains 1 atrioturbinal, 1 maxilloturbinal, 1 nasoturbinal, and at least 3 ethmoturbinals. Due to the strong development of the alveoli of the incisors, the maxilloturbinale is flected in the caviomorphs. The epiphanial foramina are present. The lamina transversalis anterior is continuous with the nasal septum so that there is a complete zona anularis in rodents. The paraseptal cartilages are continuous with the lamina transversalis anterior but not with the lamina transversalis posterior.(ABSTRACT TRUNCATED AT 400 WORDS)