The effects of habitat fragmentation on the social kin structure and mating system of the agile antechinus, Antechinus agilis

Habitat fragmentation is one of the major contributors to the loss of biodiversity worldwide. However, relatively little is known about its more immediate impacts on within-patch population processes such as social structure and mating systems, whose alteration may play an important role in extinction risk. We investigated the impacts of habitat fragmentation due to the establishment of an exotic softwood plantation on the social kin structure and breeding system of the Australian marsupial carnivore, Antechinus agilis. Restricted dispersal by males in fragmented habitat resulted in elevated relatedness among potential mates in populations in fragments, potentially increasing the risk of inbreeding. Antechinus agilis nests communally in tree hollows; these nests are important points for social contact between males and females in the mating season. In response to elevated relatedness among potential mates in fragmented habitat, A. agilis significantly avoided sharing nests with opposite-sex relatives in large fragment sites (but not in small ones, possibly due to limited nest locations and small population sizes). Because opposite-sex individuals shared nests randomly with respect to relatedness in unfragmented habitat, we interpreted the phenomenon in fragmented habitat as a precursor to inbreeding avoidance via mate choice. Despite evidence that female A. agilis at high inbreeding risk selected relatively unrelated mates, there was no overall increased avoidance of related mates by females in fragmented habitats compared to unfragmented habitats. Simulations indicated that only dispersal, and not nonrandom mating, contributed to inbreeding avoidance in either habitat context. However, habitat fragmentation did influence the mating system in that the degree of multiple paternity was reduced due to the reduction in population sizes and population connectivity. This, in turn, reduced the number of males available to females in the breeding season. This suggests that in addition to the obvious impacts of reduced recruitment, patch recolonization and increased genetic drift, the isolation of populations in habitat patches may cause changes in breeding behaviour that contribute to the negative impacts of habitat fragmentation.     

Seasonal changes in glomerular filtration rate in Antechinus stuartii (Marsupialia: Dasyuridae)

The small marsupial Antechinus stuartii experiences a synchronised life cycle that culminates in complete male mortality (within 3 weeks) following the 1 week mating period in mid-August (late winter). There are pronounced physiological changes in male A. stuartii over the life cycle and renal function was assessed for correlation with these changes. Glomerular filtration rate and urine and plasma electrolytes were determined in male and female A. stuartii in February, May, July and August. Females showed little change in glomerular filtration rate, except for pre-mating values in August which decreased. In contrast, glomerular filtration rate of males decreased significantly in July and August. Plasma sodium and chloride levels were higher in males than females and were higher in animals in July and August than in February and May. Plasma potassium levels dropped in both males and females in July and August. Plasma osmolality was higher in animals in February compared to animals from May and August. However, there were no significant sex or seasonal differences in urine electrolytes, although urea concentration was higher in females than males. Urine osmolality was higher in both sexes in July and August. There were no significant differences in total excretory rates of sodium, potassium or chloride between sexes or between seasons. Many of the alterations in renal function are correlated with known physiological and hormonal profiles in A. stuartii. This is the first observation of seasonal changes in glomerular filtration rate that are unrelated to dietary and water stresses. Accepted: 8 September 1997     

Sperm transport and storage in the agile antechinus (Antechinus agilis).

This study was an examination of the timing of ejaculation and the dynamics of sperm transport in the female reproductive tract of the agile Antechinus (Antechinus agilis) and the relationship of these parameters to single and multiple matings. Mating in this species is characteristically long compared with that of other mammals, lasting for up to 8-12 h during which time the pair remain locked together. After the first hour of mating, only approximately 40% of males had ejaculated, but by the third hour all males had ejaculated. The total number of spermatozoa extracted from the female tract remained at approximately 30 x 10(3) spermatozoa per side over the next 9 h of copulation. After completion of male/female access (12 h), approximately 56% of spermatozoa extracted were located in the lower isthmic region of the oviduct where specialized sperm storage crypts are located. The number of spermatozoa extracted from the female reproductive tract did not decline over the next 3 days, but there was a change in the distribution of spermatozoa with an increase in the proportion of extracted spermatozoa stored in the lower isthmus (approximately 76%). However, 7 to 14 days after mating, only approximately 30% of the stored spermatozoa ( approximately 9.4 x 10(3) spermatozoa per side) were still present in the isthmus. When females were mated with a second male on a consecutive day, the sperm numbers extracted from the tract were about twice that deposited during single mating, with sperm transport to the lower isthmus occurring over a similar time frame. Although the occurrence of extended copulations in the wild has not yet been confirmed, these laboratory results suggest that similar periods of copulation are likely, since completion of the ejaculation process requires at least 3 h. The extended copulation in A. agilis reduces the possibility of an early second mating, which might interfere with the normal transport and crypt colonization of spermatozoa through competition.     

Factors influencing paternity success in Antechinus agilis: last‐male sperm precedence,timing of mating and genetic compatibility

We describe the patterns of paternity success from laboratory mating experiments conducted in Antechinus agilis, a small size dimorphic carnivorous marsupial (males are larger than females). A previous study found last‐male sperm precedence in this species, but they were unable to sample complete litters, and did not take male size and relatedness into account. We tested whether last‐male sperm precedence regardless of male size still holds for complete litters. We explored the relationship between male mating order, male size, timing of mating and relatedness on paternity success. Females were mated with two males of different size with either the large or the small male first, with 1 day rest between the matings. Matings continued for 6 h. In these controlled conditions male size did not have a strong effect on paternity success, but mating order did. Males mating second sired 69.5% of the offspring. Within first mated males, males that mated closer to ovulation sired more offspring. To a lesser degree, variation appeared also to be caused by differences in genetic compatibility of the female and the male, where high levels of allele‐sharing resulted in lower paternity success.     

Acoustic features contributing to the individuality of wild agile gibbon (Hylobates agilis agilis) songs

We examined acoustic individuality in wild agile gibbon Hylobates agilis agilis and determined the acoustic variables that contribute to individual discrimination using multivariate analyses. We recorded 125 female-specific songs (great calls) from six groups in west Sumatra and measured 58 acoustic variables for each great call. We performed principal component analysis to summarize the 58 variables into six acoustic principal components (PCs). Generally, each PC corresponded to a part of the great call. Significant individual differences were found across six individual gibbons in each of the six PCs. Moreover, strong acoustic individuality was found in the introductory and climax parts of the great call. In contrast, the terminal part contributed little to individual identification. Discriminant analysis showed that these PCs contributed to individual discrimination with high repeatability. Although we cannot conclude that agile gibbon use these acoustic components for individual discrimination, they are potential candidates for individual recognition.     

Mating behaviour in Moina brachiata (Jurine, 1820) (Crustacea,Anomopoda)

Information on mating behaviour in Anomopoda isavailable for very few species only, though matelocation and recognition certainly play an importantrole in maintaining reproductive isolation betweenspecies. Ephippial females and males of Moinabrachiata were observed in a drop of water under amicroscope for 10–20 minutes. Different combinationsof specimens were used, but copulation was onlyobserved when two males and two ephippial females wereplaced together. Males were very active, and oftentried to grasp a female, whereas females usually madeattempts to escape during the entire period of mating.Three phases were recognized: capture, positioning andcopulation. The male captured the female from thedorsal side, then moved to the ventral side and tooka position with its length axis being perpendicular tothat of the female, forming a sort of cross.Thereafter the pair started rotating around the lengthaxis of the female, while the male pushed thepostabdomen into the females brood pouch. Copulationlasted from 16 to 25 seconds. When different kinds offemales were used, males showed preference forephippial females with an empty ephippium and enlargedovaries. Our results indicate that not only visual andtactile cues may be important in identifying speciesidentity and receptivity of the female, but alsochemical signals.     

The effects of habitat fragmentation via forestry plantation establishment on spatial genotypic structure in the small marsupial carnivore, Antechinus agilis

Dispersal is an important influence on species' distributions, patch colonization and population persistence in fragmented habitat. We studied the impacts of habitat fragmentation resulting from establishment of an exotic pine plantation on dispersal of the marsupial carnivore, Antechinus agilis. We applied spatial analyses of individual multilocus microsatellite genotypes and mitochondrial haplotypes to study patterns of gene flow in fragmented habitat and natural habitat 'control' areas, and how this is affected by the spatial dispersion of habitat patches, the presence of corridors and a 'mainland' source of migrants. Spatial analysis of molecular variance and partial Mantel tests confirmed the absence of cryptic barriers to gene flow in continuous habitat, which if present would confound the comparison of genetic structures in fragmented vs. unfragmented habitats. Spatial genotypic structure suggested that although dispersal was male-biased in both habitat types, fragmentation restricted dispersal of males more than that of females and the degree of restriction of male dispersal was dependent on the geographical isolation of the patch. The scale of positive genotypic structure in fragmented habitat was restricted to the two closest patches for females and the three closest patches for males. Our results provide evidence for significantly increased gene flow through habitat corridors relative to that across the matrix and for significantly lower gene flow between 'mainland' unfragmented habitat and habitat patches relative to that within either habitat type, suggesting a behavioural barrier to crossing habitat interfaces.     

Copulation behaviour in mammals: evidence that sperm competition is widespread

We review information on copulation behaviour and sperm competition in mammals using data primarily from the literature. Female mammals of many species regularly copulate with more than one male during each oestrous period. Such multi-male copulations are reported more often in social, compared with solitary species. In addition, the mates of males of polygynous species experience multi-male copulations as often as the mates of males or monogamous species. Male mammals attempt to increase their certainty of paternity through a number of reproductive tactics. Copulation frequency is higher in specks with multi-male copulations compared with other species. Consortships, which can br regarded as a form of mate guarding, are often absent from species in which more than one male copulates with each female during her oestrous period. Most solitary burrow-living small mammal species copulate in the open, whereas social species often copulate inside their burrows. Insurance copulations may occur in many species since high copulation rates occur in four circumstances: (i) when mates are reunited, (ii) when a new male takes over a female, (iii) when a strange male steals a copulation, and (iv) when an audience of males is present.     

Linstowinema breve n. sp. (Nematoda: Seuratidae) a parasite of Antechinus agilis (Marsupialia: Dasyuridae) from Australia

A new echinonematine nematode, Linstowinema breve sp. n., from the small intestine of the dasyurid marsupial Anthechinus agilis is described. The species is distinguished from its congeners by the possession of the following suite of characters: small size; first and third row of cephalic hooks similar in size; second row larger; 13-15 rows of body hooks without undulating edges on the dilated cuticle of the oesophageal region; oesophagus terminating at the level of the 5th-7th row of body hooks; ten pairs of caudal papillae; a large pair of lateral ad-cloacal papillae extend into small lateral alae. Linstowinema larvae previously recorded from A. agilis may be the same species. A key to species of the genus linstowinema is provided.     

Mating Behavior of Southern Stingrays, Dasyatis americana (Dasyatidae)

We document in detail the first complete sequence of mating events in the southern stingray, Dasyatis americana, based on observations of four matings (five separate copulations) at Grand Cayman, British West Indies and Bimini, Bahamas. These observations are significant because almost nothing is known about this important aspect of the life cycle of batoids, due to the rarity of encounters with mating animals in natural settings. Similar to mating behavior described in the manta ray, Manta birostris, the mating sequence of D. americana can be characterized as (1) 'close-following', (2) 'pre-copulatory biting', (3) 'insertion/copulation', (4) 'resting' and (5) 'separation'. Additional information gained from these observations includes the fact that (1) two of the females were mated very shortly (i.e. within minutes–hours) after parturition and (2) one of the mating events involved a single female that copulated (unforced) with two males in rapid succession. The latter observation suggests that polyandry and multiple paternity may be elements of the mating system of D. americana.     

Seasonal changes in the reproductive anatomy of male Antechinus stuartii (Marsupialia: Dasyuridae)

Male Antechinus stuartii (Marsupialia: Dasyuridae) undergo significant endocrinological changes during their life history. The endocrine changes are associated with complete male mortality. The accessory reproductive tract of males is important for seminal plasma production, and changes associated with the reproductive cycle were not described in detail by earlier studies. The present study sought to describe some of the seasonal changes in structure of the male accessory reproductive tract in relation to the known hormonal changes. The epididymis, prostate, and bulbourethral glands are relatively undifferentiated in February and May, a time when plasma concentrations of testosterone are known to be low. By July, considerable hypertrophy and differentiation of the accessory reproductive tract are observed. This is most obvious in the bulbourethral glands, which change from being indistinguishable from one another in February and May to being three large morphologically and histologically distinct glands in July. The hypertrophy and secretory activity continue into August, the breeding season. These findings correlate with the hormonal profiles found in other studies of A. stuartii. J Morphol 231:261–275, 1997. © 1997 Wiley-Liss, Inc.     

Paternity success and the direction of sexual selection in a field population of a semelparous marsupial,Antechinus agilis

Antechinus agilis is a small sexually size dimorphic marsupial with a brief annual mating period of 2-3 weeks. All males die after this period, and females give birth to up to 10 young. Mating is thought to be promiscuous, however, there is no field data to confirm this. Using microsatellites, we investigated paternity patterns over two seasons in a wild population. Male weight was significantly positively related to the number of females fertilized and with the number of offspring sired, in both years. Furthermore, selection gradients indicated selection for larger males. Both results suggest that size dimorphism in A. agilis can be explained by sexual selection for larger males. The proportion of offspring sired within litters, did not relate to male size. Therefore, larger males are more successful through higher mating access, not through their sperm outcompeting that of smaller males. As expected from their known ranging behaviour, the number of offspring within litters left unassigned to a father did not depend on the grid location of the mother. Female size did not differ between successful reproducing and unsuccessful females. However, females that weaned offspring had larger heads than females that did not wean offspring. Males did not 'prefer' mating with larger females, nor did assortative mating occur. From our results, the mating system of A. agilis is clearly promiscuous. Selection for larger males occurred in both years, even though in one year the operational sex ratio was highly female biased, suggesting that the potential reproductive rate is a better predictor of the direction of sexual selection in A. agilis.     

Social and foraging components of the home range in Antechinus stuartii (Dasyuridae: Marsupialia)

The home range of Antechinus stuartii has two components. A small area (0.94 ha for males, 0.38 ha for females) is used for foraging, while a much greater social range (mean for males in a given year may exeed 5 ha; the mean for females never exceeded 3 ha) encompasses communal nests, which are used by both males and females. The foraging range of females is stable throughout the year. The foraging range may not coincide at all, or only partially coincide with her social range. By contrast, male foraging ranges drift through the year, and generally lie within their social range. Intersexual differences reflect the different selection pressures on the sexes. Females require access to resources to sustain a costly lactation, while males require a knowledge of many possible sites for mating and early resolution of agonistic interactions with other males with which they cohabit during mating.     

Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca:Prosobranchia)

The ability of males of Littorina littorea and L. saxatilis to discriminate between mates of different sex, species and size was examined. In partner choice experiments males of L. littorea had the possibility to initiate a copulation with either a female or a male. The males did not show a preference for either sex. There was therefore no evidence that they could determine the sex of a conspecific prior to copulation. The duration of intrasexual copulation was considerably shorter than for intersexual copulation, both in the field and in laboratory experiments. For the two species, intersexual copulations were far more frequent than intrasexual ones. This can partly be explained by the difference in copulation time.Few interspecific copulating pairs were found on the shore. This may reflect a low interspecific encounter rate rather than a mechanism of species recognition. On all of these occasions, however, the active male was of L. saxatilis. It is argued that selection against precopulatory species and sex recognition is a more likely explanation than an hypothesis that states that the required mutations for precopulatory mate identification has not yet occurred. L. littorea males copulated longer with large than with small females. Copulation time was short with parasitized females, which are sterile or of low fecundity. The allocation of mating effort by males is discussed.     

Variable Female Mating Positions and Offspring Sex Ratio in the Spider Pityohyphantes phrygianus (Araneae: Linyphiidae)

Chromosomal sex determination and male heterogamety have been thought to seriously impede direct sex ratio control. However, in Pityohyphantes phrygianus, a solitary sheetweb spider with a skewed sex ratio, earlier experimental studies suggested that there are options for female control of offspring sex ratio, if females change their position during the normal mating sequence. Here we show that under natural conditions there is considerable between-female variation in positions, especially after termination of mating. Computer simulations of the orientation of female inner genitalia suggest that sperm are placed in different storage sites depending on the positions adopted. This means that a specific position after mating might potentially influence offspring sex ratio. The variance in offspring sex ratio among females in earlier experiments was binomially distributed, which leads us to conclude that females control the mean sex ratio but do not exercise direct control of the sex of individual offspring.     

Individuality in the songs of wild agile gibbons (Hylobates agilis) of Peninsular Malaysia

Recordings were made and analyzed of the female dominated duet songs by eight adult pairs of wild agile gibbons from two sites in Peninsular Malaysia. A statistical analysis of their songs revealed that individuality occurred throughout the females' songs from both sites. Individuality in gibbon songs may allow singing individuals to more efficiently and effectively locate and identify neighboring nonmate conspecifics.     

Habitat selection by the small dasyurid Ningaui yvonneae (Marsupialia: Dasyuridae) in South Australia

This study investigated the habitat preferences of Ningaui yvonneae, a small, nocturnal insectivorous marsupial of the semi‐arid regions of southern Australia. Habitat composition was assessed at both the local (trapping grid) and trap scale. Multiple linear regression revealed a significant positive relationship between captures of N. yvonneae and the abundance of hummock grass (Triodia irritans) at the local scale. At the trap scale, the interaction between Triodia irritans and shrubs was found to be significant. This is thought to reflect the importance of the availability of cover to ningauis. Triodia was an essential component of that, probably because it affords greater protection from predators. This study provides quantitative habitat data that supports the contention of other studies that N. yvonneae has a strong positive association with Triodia spp.     

The effect of female mating history on sperm precedence in the two-spot ladybird, Adalia bipunctata (Coleoptera, Coccinellidae)

Effects of two different mating regimes on sperm precedencein the two-spot ladybird, Adalia bipunctata, were studied usingthe polymorphic gene for melanism as a marker for paternity.Virgin nonmelanic females (homozygous recessive) were matedto nonmelanic male(s) and then, after laying fertilized eggs,were mated to a melanic male of known genotype. The resultsafter the two successive single matings showed a highly variabledegree of paternity of the second male. Initial multiple matingwith nonmelanic males did not alter the pattern of paternityafter the subsequent single mating with a melanic male, butit had two other effects: (1) the female showed an increasein rejection behavior, and (2) a longer copulation was requiredfor high success of the melanic male. Additional observationsin which families were reared from beetles collected in copulain the field demonstrated that sperm competition also occursunder natural conditions. The outcome of the competition wasvariable with frequent sperm mixing.