Malurus Coronatus Research Articles

The population genetics of the western purple-crowned fairy-wren (Malurus coronatus coronatus), a declining riparian passerine

We investigate the population genetic structure of the declining western subspecies of the purple-crowned fairy-wren (Malurus coronatus coronatus) in order to guide conservation management recommendations for this riparian habitat specialist. Our analysis of multilocus microsatellite data, from 79 individuals sampled from across the species’ range, indicates that M. c. coronatus occurs as genetically differentiated subpopulations that correspond to catchment boundaries or expansive gaps in habitat along waterways. The genetic similarity of large populations of fairy-wrens on four catchments (Fitzroy, Durack, Drysdale and Victoria) indicates widespread recent gene flow, whereas the high genetic distinctiveness of the Bindoola and Isdell catchments may reflect the current geographic isolation of these smaller populations. Genetic differentiation of these smaller geographically isolated populations affirms the negative effect that habitat degradation and fragmentation can have on population connectivity. A regional-scale approach to conservation with a focus on preventing degradation and enhancing connectivity may be critical to safeguard the persistence of M. c. coronatus subpopulations.

Habitat geometry does not affect levels of extrapair paternity in an extremely unfaithful fairy-wren

Density of potential mates has often been proposed to explain the enormous variation in extrapair paternity. However, density is often confounded by other ecological factors that might affect extrapair paternity in their own way. Furthermore, extrapair mating shows strong phylogenetic inertia, making both meaningful intra- and interspecific comparisons difficult. An extreme way to change density is through habitat fragmentation that reduces connectivity between territories. Recently, habitat connectivity was hypothesized to explain the surprising discovery of a virtually monogamous species among the world’s most unfaithful bird genus. The monogamous Malurus coronatus lives in narrow riparian strips that limit contact with neighbors to both extreme ends of territories, whereas Malurus species with high levels of extragroup paternity typically live in high-connected habitat in which they are surrounded by neighbors. Here, we test the habitat geometry hypothesis by comparing levels of extragroup paternity of Malurus elegans living in fragmented low-connected habitat and in high-connected habitat. We found that M. elegans does not have lower levels of extragroup paternity in low-connected habitat (68%) than in high-connected habitat (56% of offspring), indicating that connectivity does not limit opportunities for extragroup paternity. Furthermore, there was no evidence that females in low-connected habitat gained extragroup paternity further away or from less sires or that they were more likely to be closely related to their social mate. We conclude that behavioral plasticity in response to density-dependent cost and benefits of mating behavior does not explain intrageneric variation in extragroup paternity in Malurus. Furthermore, habitat fragmentation may not strongly affect inbreeding risk in this species.

Breeding synchronization facilitates extrapair mating for inbreeding avoidance

Extrapair (EP) mating can enable females to reduce the negative effects of inbreeding. However, opportunities for EP mating are often ecologically or demographically constrained, and it is unclear whether and how females can overcome these constraints. Here, we show that fitness costs from inbreeding in purple-crowned fairy-wrens (Malurus coronatus) are severe (≥30% reduction in hatchability of eggs in incestuous pairs). Although EP paternity was low for unrelated pairs (3% of broods), and negatively affected by low breeding density, 46% of broods produced by incestuously paired females contained EP young that were almost always sired by a direct neighbor. Moreover, females in incestuous pairs initiated egg laying more synchronously with direct neighbors than females in an unrelated pair, and more synchronous broods were more likely to contain EP young. Because male sperm stores are highest around their own partner’s fertile period, our results imply that incestuously paired females achieve high levels of EP paternity by adjusting the timing of egg laying to match peak fertility of neighboring males. This example of a dramatic increase in EP paternity due to subtle behavioral adjustments illustrates that negative effects of inbreeding exert strong selection for female strategies to facilitate EP mating, and this may contribute to unexplained variation in EP paternity.

Conservation of the Patchily Distributed and Declining Purple-Crowned Fairy-Wren (Malurus coronatus coronatus) across a Vast Landscape: The Need for a Collaborative Landscape-Scale Approach

Conservation of species that are patchily distributed must consider processes that influence both the occurrence of individuals within patches, and the persistence of populations across multiple habitat patches within the landscape. Here we present a rare regional assessment of the population size and distribution of a patchily distributed, threatened species, the purple-crowned fairy-wren (Malurus coronatus coronatus), across a vast landscape. We used data from aerial vegetation mapping of waterways, with on-ground bird surveys to predict the occurrence of suitable habitat for M. c. coronatus across 14 catchments in the Kimberley region of Western Australia. Suitable habitat was extremely limited (305 km of riparian vegetation) and fragmented (342 patches) along the 2700 km of waterway surveyed within catchments where the species occurs. Populations were predicted to be large on the Fitzroy, Durack and Drysdale catchments, and small on the Isdell and northern Pentecost catchments, and a total population of 2834 to 4878 individuals could be supported. The sub-populations spanned numerous patches of habitat across multiple properties of varying tenure. Therefore, a landscape-scale approach to conservation management, across multiple tenures, is critical to safe-guard connectivity within populations. The greatest benefit may be achieved by a combination of broad-scale actions to reduce the impact of ubiquitous threatening processes, and fine-scale targeted effort in areas where populations are most vulnerable. Controlling access of stock to waterways and management of fire are most important to conserve suitable habitat. Such a landscape-scale approach to conservation may be of benefit to other patchily distributed species.

Male Songbird Indicates Body Size with Low-Pitched Advertising Songs

Body size is a key sexually selected trait in many animal species. If size imposes a physical limit on the production of loud low-frequency sounds, then low-pitched vocalisations could act as reliable signals of body size. However, the central prediction of this hypothesis – that the pitch of vocalisations decreases with size among competing individuals – has limited support in songbirds. One reason could be that only the lowest-frequency components of vocalisations are constrained, and this may go unnoticed when vocal ranges are large. Additionally, the constraint may only be apparent in contexts when individuals are indeed advertising their size. Here we explicitly consider signal diversity and performance limits to demonstrate that body size limits song frequency in an advertising context in a songbird. We show that in purple-crowned fairy-wrens, Malurus coronatus coronatus, larger males sing lower-pitched low-frequency advertising songs. The lower frequency bound of all advertising song types also has a significant negative relationship with body size. However, the average frequency of all their advertising songs is unrelated to body size. This comparison of different approaches to the analysis demonstrates how a negative relationship between body size and song frequency can be obscured by failing to consider signal design and the concept of performance limits. Since these considerations will be important in any complex communication system, our results imply that body size constraints on low-frequency vocalisations could be more widespread than is currently recognised.

Causes of Ring-Related Leg Injuries in Birds – Evidence and Recommendations from Four Field Studies

One of the main techniques for recognizing individuals in avian field research is marking birds with plastic and metal leg rings. However, in some species individuals may react negatively to rings, causing leg injuries and, in extreme cases, the loss of a foot or limb. Here, we report problems that arise from ringing and illustrate solutions based on field data from Brown Thornbills (Acanthiza pusilla) (2 populations), Siberian Jays (Perisoreus infaustus) and Purple-crowned Fairy-wrens (Malurus coronatus). We encountered three problems caused by plastic rings: inflammations triggered by material accumulating under the ring (Purple-crowned Fairy-wrens), contact inflammations as a consequence of plastic rings touching the foot or tibio-tarsal joint (Brown Thornbills), and toes or the foot getting trapped in partly unwrapped flat-band colour rings (Siberian Jays). Metal rings caused two problems: the edges of aluminium rings bent inwards if mounted on top of each other (Brown Thornbills), and too small a ring size led to inflammation (Purple-crowned Fairy-wrens). We overcame these problems by changing the ringing technique (using different ring types or larger rings), or using different adhesive. Additionally, we developed and tested a novel, simple technique of gluing plastic rings onto metal rings in Brown Thornbills. A review of studies reporting ring injuries (N = 23) showed that small birds (<55 g body weight) are more prone to leg infections while larger birds (>35 g) tend to get rings stuck over their feet. We give methodological advice on how these problems can be avoided, and suggest a ringing hazard index to compare the impact of ringing in terms of injury on different bird species. Finally, to facilitate improvements in ringing techniques, we encourage online deposition of information regarding ringing injuries of birds at a website hosted by the European Union for Bird Ringing (EURING).

Sperm storage reflects within- and extra-pair mating opportunities in a cooperatively breeding bird

In passerine birds, storage and maturation of sperm takes place in the cloacal protuberance (CP), an external swelling of the reproductive organ. The considerable variation in CP size among species is presumed to be a consequence of varying levels of sperm-competition, but whether individual variation in CP size within a species also reflects sperm competition is not well established. Here, we study temporal variation in male CP size in relation to within-pair and extra-pair mating opportunities and cuckoldry risk in purple-crowned fairy-wrens Malurus coronatus. This is a socially monogamous cooperatively breeding passerine that can breed year-round and has low levels of extra-pair paternity (in 6 % of broods). We show that male CP size sharply increased a few weeks before, and rapidly regressed after his partner laid eggs, consistent with a cost of its maintenance and/or sperm production. Surprisingly, despite low levels of extra-pair paternity, CP size of non-breeding and pre-breeding males was positively correlated with the number of breeding females in the population, suggesting that CP size is sensitive to extra-pair mating opportunities. However, CP sizes do not seem to reflect cuckoldry risk: CP size of dominant males was unaffected by the presence of a subordinate that was unrelated to the dominant female, although those subordinates occasionally sire offspring, and had a larger CP than subordinates living with their mother. Our results suggest that, even in a species with low levels of extra-pair paternity, individual investment in sperm storage reflects both within-pair and, albeit to lesser extent, extra-pair mating opportunities.

Inferring population connectivity across the range of the purple-crowned fairy-wren (Malurus coronatus) from mitochondrial DNA and morphology: implications for conservation management

Knowledge of population structure and patterns of connectivity is required to implement effective conservation measures for the purple-crowned fairy-wren (Malurus coronatus), a threatened endemic of northern Australia. This study aimed to identify barriers to dispersal across the distribution of M. coronatus, investigate the impact that the recent declines may have on population connectivity, and propose conservation actions to maintain natural patterns of gene flow. Analysis of mitochondrial DNA sequences from 87 M. coronatus identified two phylogenetic clusters that corresponded with the phenotypically defined western (M. c. coronatus) and eastern (M. c. macgillivrayi) subspecies. The genetic divergence between these subspecies was consistent with isolation by a natural barrier to gene flow, and supports their separate conservation management. Within the declining M. c. coronatus, the lack of genetic divergence and only slight morphological difference between remnant populations indicates that populations were recently linked by gene flow. It is likely that widespread habitat degradation and the recent extirpation of M. c. coronatus from the Ord River will disrupt connectivity between, and dynamics within, remnant populations. To prevent further declines, conservation of M. coronatus must preserve areas of quality habitat and restore connectivity between isolated populations.

Influence of fine‐scale habitat requirements and riparian degradation on the distribution of the purple‐crowned fairy‐wren (Malurus coronatus coronatus) in northern Australia

AbstractSpecies distributions are influenced by variation in environmental conditions across many scales. Knowledge of fine‐scale habitat requirements is important for predicting species occurrence and identifying suitable habitat for target species. Here we investigate the perplexing distribution of a riparian habitat specialist, the western subspecies of the purple‐crowned fairy‐wren (Malurus coronatus coronatus), in relation to fine‐scale habitat associations and patterns of riparian degradation. Surveys of vegetation attributes, river structure and disturbance indicators that are likely to be causal determinants of the species occurrence were undertaken at 635 sites across 14 catchments. Generalized Linear Mixed Modelling demonstrated that the probability of purple‐crowned fairy‐wren occurrence increased with Pandanus aquaticus crown cover, shrub density and height of emergent trees, while riparian structure and signs of cattle were indirect predictors of occurrence. As our study area predominantly contained Pandanus type habitat, we failed to identify river grass as an important component of habitat. Predictions from a cross‐validated model of purple‐crowned fairy‐wren occurrence suggested distribution is constrained by three factors: (i) low quality of local habitat within catchments where the species occurs; (ii) broad‐scale reduction in habitat quality that has resulted in extinction of the species from parts of its range; and (iii) unmeasured variables that limit the exploitation of suitable habitat. The reliance of the species on dense shrubby understorey suggests conservation efforts should aim to maintain the complexity of understorey structure by managing fire and grazing intensity. Efforts to halt the continuing decline of riparian condition and maintain connectivity between areas of quality habitat will help to ensure persistence of riparian habitat specialists in northern Australia.

Multiple Benefits Drive Helping Behavior in a Cooperatively Breeding Bird: An Integrated Analysis

Several hypotheses exist to explain the seemingly altruistic helping behavior of cooperative breeders, although the general utility of these hypotheses remains unclear. While the potential importance of inclusive fitness benefits (kin selection) is traditionally widely appreciated, it is increasingly recognized that direct benefits may be more important than assumed. We use an integrative two-step framework to assess support for current hypotheses in purple-crowned fairy wrens, a species where subordinates vary in relatedness to breeders and helping increases productivity. After establishing that assumptions of pay-to-stay and social prestige hypotheses (predicting that helping functions as "paying rent" to stay on the territory or as a signal of individual quality, respectively) were not met and that parentage by subordinates is extremely rare, we tested whether subordinates adjusted nestling feeding rates following the predictions of the kin selection and group augmentation hypotheses. Benefits of kin selection result from investment in relatives, and group augmentation benefits accrue when subordinates invest more in their own future helpers, for example, when they have a better chance of inheriting the breeding position. We found that subordinates fed siblings more than unrelated nestlings, indicating that kin selection could facilitate cooperation. Moreover, the effect of relatedness on feeding effort varied depending on the probability of inheriting a breeding position, suggesting that active group augmentation can explain investment by unrelated subordinates. This statistical interaction would have gone undetected had we not considered both factors simultaneously, illustrating that a focus on single hypotheses could lead to underestimation of their importance in explaining cooperative breeding.

No evidence for offspring sex-ratio adjustment to social or environmental conditions in cooperatively breeding purple-crowned fairy-wrens

When fitness returns or production costs vary between male and female offspring, selection is expected to favor females that adjust offspring sex ratio accordingly. However, to what extent vertebrates can do so is the subject of ongoing debate. Here, we explore primary sex ratios in 125 broods of cooperatively breeding purple-crowned fairy-wrens Malurus coronatus. We expected that females might adjust offspring sex ratio because this passerine species experiences considerable variation in social and environmental conditions. (1) However, although helpers substantially increase parental fitness, females (particularly in pairs and small groups) did not overproduce philopatric males (helper-repayment hypothesis). (2) Sex-ratio adjustment based on competition among individuals (helper-competition hypothesis) did not conceal helper-repayment effects or drive sex allocation on its own: while high-quality territories can accommodate more birds, brood sex ratios were independent of territory quality, alone or in interaction with group size. (3) Additionally, males are larger than females and are possibly more costly to produce (costly sex hypothesis), and (4) female offspring may benefit more from long-term effects of favorable conditions early in life (Trivers–Willard hypothesis). Nonetheless, large seasonal variation in food abundance was not associated with a consistent skew in primary sex ratios. Thus, overall, our results did not support the main hypotheses of adaptive sex-ratio adjustment in M. coronatus. We discuss that long-term differential costs and benefits may be insufficient to drive evolution of primary sex-ratio manipulation by M. coronatus females. More investigation is therefore needed to determine the general required sex differences in long-term fitness returns for mechanisms of primary sex-ratio manipulation to evolve.

The distribution and status of the western subspecies of the Purple-crowned Fairy-wren (Malurus coronatus coronatus)

Knowledge of the distribution of a species, although fundamental for directing conservation, is often lacking for rare or declining species or those occurring in remote locations. The western subspecies of the Purple-crowned Fairy-wren (Malurus coronatus coronatus) is a riparian habitat specialist of the Kimberley and Victoria River regions of northern Australia. Although listed as vulnerable nationally, there is little information on its current distribution and population trends. We conducted a comprehensive survey across 14 river catchments within the potential distribution of the subspecies, and identified changes in distribution by comparing the results of the survey with historical records. Western Purple-crowned Fairy-wrens were recorded on the Fitzroy, Isdell, Durack, Pentecost, Drysdale and Victoria River systems. The species was not detected in the Ord or the Sale systems despite previous reports, nor was it found in the Carson, Calder, Meda, Forrest, Berkeley or King George catchments. We suggest these last three catchments were wrongly included in previous estimates of the distribution of the species, when in fact Purple-crowned Fairy-wrens were potentially never there. We furthermore identify the populations of Purple-crowned Fairy-wrens most in need of conservation intervention and recommend excluding cattle to prevent further habitat degradation and population declines. These results are a crucial baseline from which to monitor future population trends of the Purple-crowned Fairy-wren and direct conservation management.

Multiple benefits of cooperative breeding in purple‐crowned fairy‐wrens: a consequence of fidelity?

1. Kin selection is one of the mechanisms that can explain apparent altruism by subordinate individuals in cooperatively breeding species, if subordinates boost the production of kin. We compared productivity and breeder survival in pairs with and without subordinates in a genetically monogamous cooperatively breeding bird, the purple-crowned fairy-wren Malurus coronatus. 2. Additive effects of subordinate help increased productivity. Total feeding rates to the nest were increased by two or more subordinates, and fledgling production was greater in larger groups. Not all subordinates contributed to nestling feeding, and the effect of group size was greater when non-contributors were excluded from analyses, suggesting that increased fledgling production was a direct result of help. 3. Compensatory effects of subordinate help improved breeder survival. Assisted breeders reduced their workload by 20-30%, irrespective of the number of helpers. Although re-nesting intervals were not affected by group size, reduced breeder feeding rates resulted in improved survival and breeders in larger groups survived better. 4. Subordinates and nestlings are usually progeny of the breeding pair in this species, and benefits of cooperative breeding are very different from three congeners with extremely high levels of extra-group paternity (EGP). In these Malurus, fledgling production and survival of male breeders are not enhanced in larger groups. This is consistent with the expectation that kin-selected benefits vary with relatedness, and thus levels of EGP. 5. We tested whether benefits of cooperative breeding in 37 avian species varied with levels of extra-group mating. Both direct and phylogenetically controlled comparisons showed that improvement of (male) breeder survival and enhanced productivity are more likely when fidelity is higher, as predicted when investment of subordinates correlates with relatedness to offspring. This pattern highlights the importance of considering the genetic mating system for understanding the evolution of cooperative breeding.

Ian Cecil Robert Rowley 1926–2009

Ian Cecil Robert Rowley 1926–2009

A Description of the Primary Habitat of the Purple-crowned Fairy-wren 'Malurus Coronatus Coronatus' in the Victoria River District, N.T

The habitat of the Purple-crowned Fairy-wren in the Victoria River District is described, with particular reference to the density of the riparian grass Chionachne cyathopoda, based on 75 vegetation transects sampled in 39 fairy-wren territories. This paper also clarifies confusion about the primary habitat of the fairy-wren with respect to another riparian grass, Mnesithea rottboellioides, which shares with C. cyathopoda the common name Canegrass. To assist successful management and conservation we provide a guide to correctly distinguish these two grasses and recommend the use of the common name Rivergrass for C. cyathopoda and Northern Canegrass for M. rottboellioides.

Radical loss of an extreme extra-pair mating system

BackgroundMating outside the pair-bond is surprisingly common in socially monogamous birds, but rates of extra-pair paternity (EPP) vary widely between species. Although differences in life-history and contemporary ecological factors may explain some interspecific variation, evolutionary forces driving extra-pair (EP) mating remain largely obscure. Also, since there is a large phylogenetic component to the frequency of EPP, evolutionary inertia may contribute substantially to observed EP mating patterns. However, the relative importance of plasticity and phylogenetic constraints on the incidence of EP mating remains largely unknown.ResultsWe here demonstrate very low levels of EPP (4.4% of offspring) in the purple-crowned fairy-wren Malurus coronatus, a member of the genus with the highest known levels of EPP in birds. In addition, we show absence of the suite of distinctive behavioral and morphological adaptations associated with EP mating that characterize other fairy-wrens. Phylogenetic parsimony implies that these characteristics were lost in one speciation event. Nonetheless, many life-history and breeding parameters that are hypothesized to drive interspecific variation in EPP are not different in the purple-crowned fairy-wren compared to its promiscuous congeners.ConclusionSuch radical loss of an extreme EP mating system with all associated adaptations from a lineage of biologically very similar species indicates that evolutionary inertia does not necessarily constrain interspecific variation in EPP. Moreover, if apparently minor interspecific differences regularly cause large differences in EPP, this may be one reason why the evolution of EP mating is still poorly understood.

Do male paternity guards ensure female fidelity in a duetting fairy-wren?

Most socially monogamous bird species engage in extra-pair mating, and consequently males may adopt various behavioral strategies to guard paternity. However, the relationship between mate guarding and extra-pair paternity is unclear: low levels of extra-pair paternity can be associated either with no mate guarding or with intense mate guarding. We investigate paternity guards in the purple-crowned fairy-wren (Malurus coronatus), a duetting species where extra-pair paternity is rare. This species is unusual in a genus known for extremely high levels of extra-pair mating. We examine the behavioral interactions between the sexes underlying these low rates of extra-pair paternity and show that male purple-crowned fairy-wrens do not use frequent copulation or courtship feeding to assure paternity or guard females acoustically by duetting. Males maintain close proximity to females both when they are fertile and when they are not breeding and do not invest in courtship displays to extra-pair females. Consistent with predictions of theoretical models, low extra-pair paternity in this species may be related to female fidelity rather than male paternity assurance strategies, but short-term removal of males would be necessary to test this experimentally.

Coordination between the sexes for territorial defence in a duetting fairy-wren

In territorial songbirds both sexes depend on possession of a territory, yet its defence is traditionally regarded as the domain of male birds. Studies on duetting species have examined both male and female roles in territorial defence, often focusing on potential reproductive conflict between the pair members. Surprisingly, interactions between defending pair members are rarely investigated. We combined observations on undisturbed pairs of purple-crowned fairy-wrens, Malurus coronatus, with experimental simulation of a strong territorial threat to quantify coordination between partners during territorial defence. Pairs formed strongly cohesive partnerships: males and females naturally spent a high proportion of time close together (75% within 5 m) in dense riparian habitat, and coordinated half their songs to form duets. Partners that spent more time close together also coordinated more of their songs to form duets. When we presented playback simulating paired intruders, females responded as aggressively as males. Intensity of approach to the playback speaker was highly correlated within pairs. Spatial coordination between partners increased dramatically when they responded to simulated intrusion so that they spent virtually all their time within 2 m of one another, and most of the songs they sang were coordinated to form duets. We conclude that the highly coordinated vocal and approach responses of male and female purple-crowned fairy-wrens to simulated intrusion in the nonbreeding season are not coincidental, or a consequence of sexual conflict over mating, but instead are likely to represent cooperative territorial defence of a valuable shared resource.

Helpers liberate female fairy-wrens from constraints on extra-pair mate choice

In cooperatively breeding superb fairy-wrens (Malurus cyaneus), all males contribute to the feeding and defence of young. Despite the expectation that such paternal care should be directed only to relatives, DNA fingerprinting revealed that most offspring (76%, 138 out of 181) were sired by extra-group males that contributed no care, and that almost all broods (95%, 38 out of 40) contained young sired by extra-group fathers. This is the highest known incidence of cuckoldry. This remarkable mating system is produced by female control of fertilization and consistent preference for certain extra-group male genotypes. This choice leads to the production of sons that also gain extra-group fertilizations. One constraint on the extra-pair mate choice of females is the level of parental assistance received from males. Males living in pairs contribute relatively more parental care and are more likely to gain paternity in one of their broods (85%, 11 out of 13) than dominant males in multi-male cooperative groups (30%, 8 out of 27). In groups helpers compensate for the lower parental assistance of dominant males, so the total feeding rate is similar between pairs and groups. This suggests that females allow males in pairs more paternity to ensure their assistance with parental care. Helpers provide an alternative source of paternal investment, and allow females to express unrestricted mate choice. Mating options for females in other species with female control of fertilization may also reflect a trade-off between acquiring the genes of high-quality males for their offspring and parental care of those offspring.

The Purple-crowned Fairy-wrenMalurus coronatus. I. History, Distribution and Present Status

The two subspecies of Purple-crowned Fairywrens Malurus coronatus coronatus (Kimberleyan) and M.c. rnacgillivrayi (Carpentarian) occupy very restricted riverside vegetation across the Wet-Dry tropics of northern Australia between the 400 mm and 1000 mm isohyets; they are separated by 200 km of unsuitable habitat. The history of settlement, land degradation and the current status of these birds is dealt with in five localities. The Kimberleyan subspecies is estimated to be below 7000 individuals, to be vulnerable and in need of conservation measures. The Carpentarian subspecies is at least twice as numerous and is not threatened at present; its future status will need monitoring because of its restricted habitat. In the past decade several new national parks that are strategically well placed for populations of M. coronatus have been gazetted. Even more importantly, land management appears to be moving out of the exploitive, habitat degrading stage towards practices that minimise erosion and maintain the riverside as a renewable resource. This should benefit the survival of the species, especially M. c. coronatus.