Molecular methods for the genetic identification of salmonid prey from Pacific harbor seal (Phoca vitulina richardsi) scat

Twenty-six stocks of Pacific salmon and trout (Oncorhynchus spp.), representing evolutionary significant units (ESU), are listed as threatened or endangered under the Endangered Species Act (ESA) and six more stocks are currently being evaluated for listing. The ecological and economic consequences of these listings are large; therefore considerable effort has been made to understand and respond to these declining populations. Until recently, Pacific harbor seals (Phoca vitulina richardsi) on the west coast increased an average of 5% to 7% per year as a result of the Marine Mammal Protection Act of 1972 (Brown and Kohlman2). Pacific salmon are seasonally important prey for harbor seals (Roffe and Mate, 1984; Olesiuk, 1993); therefore quantifying and understanding the interaction between these two protected species is important for Morphobiologically sound management strategies. Because some Pacific salmonid species in a given area may be threatened or endangered, while others are relatively abundant, it is important to distinguish the species of salmonid upon which the harbor seals are preying. This study takes the first step in understanding these interactions by using molecular genetic tools for species-level identification of salmonid skeletal remains recovered from Pacific harbor seal scats.

Impact of community-based fisheries management practices on production and species diversity in Turag-Bangshi basin floodplains in Kaliakoir, Gazipur, Bangladesh

The study was conducted to investigate the communities perception and compliance to community-based fisheries management (CBFM) in Turag-Bangshi floodplains under Kaliakoir, Gazipur District. Measures such as ban on use of the harmful fishing gears, seasonal fishing closure, halt of fry fishing, halt of dewatering of beels and the impact of establishment of sanctuaries on fish production and species diversity were introduced by MACH project. Almost all members of the communities in Turag-Bangshi MACH (Management of Aquatic Ecosystem through Community Husbandry) site welcomed the introduction and complied with the implementation of all management measures which helped stopped use of harmful fishing gears, ensured survival and breeding of brood fish in the rainy season, protected and allowed fry to grow big, restored lost and degraded fisheries and organized communities for sustainable development of the fisheries. A total of 51 species of fishes were found in Makosh beel (natural depression). Among these, small indigenous species (SIS) under Cyprinidae family (Puntius sophore) was the most dominant. Many species available in the past recorded disappeared from the Makosh beel due to loss of habitat and industrial pollution that damaged spawning and nursery grounds of fish. Introduction of some selective native endangered species (Nandus nandus, Notopterus notopterus, Ompok pabda and Labeo calbasu) by MACH in the Turag-Bangshi water bodies increased diversity of species from 82 to 95. Over a period of five years during MACH intervention, the average production remained nearly 200% higher than the baseline production of 57 kg/ha to present 207 kg/ha due to maintaining sanctuaries and the closed fishing seasons. Per capita daily fish consumption of the surrounding communities also increased by 78% (from 27 to 48 g/person/day) which is much higher than the national average fish consumption in Bangladesh. The implementation of community-based MACH project management measures substantially improved fish habitat, production, consumption and socio-economic conditions of the surrounding communities. The model can be used to improve the floodplains of Bangladesh.

Marketing of small indigenous species of fish (SIS) in three selected fish markets of Mymensingh

A socio-economic survey was conducted round the year in three fish markets at Mymensingh, Bangladesh. The selected markets were categorized as rural market (Sutiakhali market), a peri-urban market (Kamal Ranjeet market, BAU) and an urban market (Notun Bazar market, Mymensingh town). It was learnt from the survey that the availability of Small Indigenous Fish Species (SIS) declined to a great extent over the last few years and at presently many of such fish species are either threatened or at the edge of extinction. The supply of SIS was highest in KR market (37% of total) and more or less similar in Notun Bazar and Sutiakhali fish market (25 and 27% respectively). The total supply of SIS fluctuated from 25% to 35% throughout the year in these markets. About 48 SIS were found in the sampled markets over the survey period. The highest number of species (45) was found in KR market followed by Notun Bazar (42) and Sutiakhali (37) fish markets. During the survey, three critically endangered species namely, schilbid catfish, garua catfish and rita were found in these markets. Beside these, other 11 and 10 species were listed to be endangered and vulnerable respectively. The biodiversity of 21 SIS found in three markets were no threat at all. Three species (guntea loach, Indian glass barb and flying barb) were 'data deficient' as reported by the IUCN Red Book (IUCN-Bangladesh 2000). From the supply point of view small prawn, spotted snakehead, stinging catfish, pool barb, striped dwarf catfish, Gangetic mystus, walking catfish and tank goby were the prominent fish. The least available species found in this survey were lesser spiny eel, barred spiny eel, Gangetic ailia, freshwater garfish, zig-zag eel, flying barb, Ganges river sprat, freshwater river shad and dwarf gourami. The weight of SIS available in Notun bazar was highest and nearly double than other two markets. There was no significant difference recorded in the supply of SIS in Sutiakhali and KR markets. The average monthly SIS supply was 185, 192 and 467 kg in KR, Sutiakhali and Notun Bazar, respectively; therefore, the cumulative average supply was 844 kg per month in three markets. The price of SIS ranged widely from taka 50-450/kg depending on species, location of market, time of purchase and the condition of fish. In general small prawn, ticto barb, dwarf gourami, Gangetic leaffish, and Annandale loach were sold at a lower price (ranged taka 50-100/kg) and these species could be considered at the bottom of the market-price list. Other SIS like walking catfish, climbing parch, butter catfish, cotio and schilbid catfish valued as highest price (ranged taka 150-450/kg). There was no specific marketing chain for SIS in Mymensingh region. The components of marketing channels and their expansion varied with seasons and locations. The general pattern, however, was as this - after buying fish from fish farmer/fishermen, middlemen (locally known as Foria) used to buy fish to wholesale market and sell to the wholesalers. The retailers used to buy fish from wholesaler through auction to the highest bidders. The retailers then send the fish to particular market where the fish reached the consumers. The livelihood strategy of SIS retailers in three fish markets showed that socio-economic constraints such as low income, poor educational background, low economic status and lack of capital are the main constrains [sic]. Most of the retailers proposed that government should control the fish price throughout the year, so that the producers can get reasonable and stable price. Construction of cold storage and preservation facilities at market sites, improvement of road and communication, improvement of physical market facilities and reduction of market chain is essential. Credit facilities, improvement of their standard of living, health and sanitary condition, housing condition, children education and access to drinking water facilities were identified as additional aspects to improve socio-economic condition of SIS retailers.

Fish species diversity in the Victoria and Kyoga lake basins: their conservation and sustainable use

Introduction of exotic fish species especially the Nile perch Lates niloticus, is believed to be responsible for the decline of fish species diversity in lakes Victoria, Kyoga and Nabugabo.About 60% of the haplochromine cichlids are thought to have become extinct from L. Victoria due to predation by the Nile perch. However there are many lakes satelite to the lakes Victoria and Kyoga basins which still have fish fauna similar to that of the main lakes. many of the satellite lakes are separated from the main lakes in, which Nile perch was introduced by extensive swamps that provide a barrier to Nile perch .A survey was carried out in a number of these satelite lakes and an inventory made of existing fish species. Their distribution and relative abundances were also determined. The lakes studied included Nawampasa, Nakuwa,Kawi Lamwa Gigate, Nyaguo, Agu, Nabugabo. Kayanja, Kaytigi, Mburo, Kachera and Wamala.Some habitats within the main lakes Victoria and Kyoga, especially those with rocky outcrops· and macrophyte cover that provide refugia for endangered species from Nile perch,were also surveyed) Various stations along the River Nile were also sampled to quantify the fish species that are still resent. Kyoga minor lakes were found to have the highest number of fish species especially of haplochromine cichlids. Many haplochromine trophic groups that were thought to be extinct from 1. Victoria still occur in these lakes.!Some of the satellite lakes, especially lakes Kayugi, Mburo and Kachera still contain .healili populations of oreochromis. I esculentus that could be used as brood stock in fish farming. Many of these lakes should .I ( I therefore be protected for conservation offish species diversity

REPORT ON THE DISTRIBUTION, POPULATION, AND ECOLOGY OF THE YUNNAN SNUB-NOSED MONKEY (RHINOPITHECUS-BIETI)

The Yunnan snub-nosed monkey (Rhinopithecus bieti), an endangered species in China, has received more protection in theory than in practice. Therefore it is on the very verge of extinction. The population of the species was estimated less than 2,000 individuals spread in 19 distinct groups. It was confirmed that the monkey was confined to the Yunling Mountain System, the area between the Yangtze River (Changjiang, aka Jinshajiang) to the east and the Mekong River (Lancangjiang) to the west. We further concluded that a lowland belt to the east, about 100 km long and 20 - 30 km wide was not suitable habitat for the monkeys, and appeared to serve as the natural ecogeologic barrier for the species. Our results indicated that the southern limit of the distribution was at Longma (26-degrees 14'N), and that the northern limit of the distribution was at Xiaochangdu (29-degrees 20'N). The distribution area of the species was substantially smaller than previously estimated. There were substantial ecological differences between the southern and northern parts of the species range. The monkey was found only in fir-larch forest.

Range-wide Yangtze freshwater dolphin expedition: The last chance to see Baiji?

Background, Aim and Scope. There are two species of fresh water cetaceans surviving in the Yangtze River system in China: Baiji (Lipotes vexillifer) and Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis). As a result of the expansion of human activities on the river, their distribution ranges appear to be decreasing and in the case of the Baiji, are even being restricted to several sections. The Baiji is the world's most critically endangered cetacean species with a population estimated at only a few tens of individuals. The Yangtze finless porpoise is the world's only freshwater-adapted population of the species, and it has been estimated that only around 1,000 individuals remain in the river system. In order to prevent the extinction of Baiji and a sharp decline in the abundance of the porpoise, in situ conservation (i.e. in the river) and two ex situ conservation (i.e. in semi-natural reserves and in captivity) strategies were proposed and have been implemented since the early 1990s. In view of both the severely endangered status of the animals and the severely degraded conditions of their habitats, the feasibility and actual status of these two strategies are raised for discussion. Main Features. The threats faced by the cetaceans are mainly from the unfettered exploitation of the river's resources. In the past 20 years, five nature reserves have been established along the river. Imposing maximum prohibition of harmful and illegal fishing methods in the reserves might prolong the process of extinction of these cetaceans in the wild, but so far, the administrative measures taken in the reserves have not yet kept the abundance from sharply declining. As human use of the river and its resources is expected to intensify for many decades into the future, the ability of the river to continue to support these species is certainly undecided. Therefore, rescuing animals from the river and establishing viable breeding populations in semi-natural reserves, in which the environment is similar to the main stream of the river, and in captivity, has to be considered urgently as the short-term goal of ex situ strategies. Since the abundance of porpoises is higher than that of the Baiji, we have first established breeding populations of them in the semi-natural reserves and in captivity. But, considering the extremely low density of Baiji in the river, an immediate range-wide Yangtze Baiji survey is an urgent need for locating and capturing sufficient Baiji for successfully establishing a breeding population of them in semi-natural reserves. Results. Two semi-natural reserves (in Shishou, Hubei Province, and Tongling, Anhui Province) have been set up along the river in order to establish breeding populations of the Baiji and the porpoises. So far, several small groups of porpoises that were caught in the main stream of the river have successively been introduced into the semi-natural reserves. Under careful management, these animals in both of the semi-natural reserves not only survive, but can also reproduce naturally and successfully. At least one or three calves were born in each reserve each year. Additionally, a breeding group of porpoises is being established at the Baiji Dolphinarium at the Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan. There are presently four adults and one calf living in the Dolphinarium. The calf, born in July, 2005, is the first captive bred Yangtze Finless Porpoise in the world. In preparation for the range-wide Yangtze cetacean survey, a 9-day pilot expedition on the river near Wuhan was conducted in March, 2006, in order to develop methods for locating the Baiji. No Baiji were expected to be seen in such a short period but about 40 porpoise sightings were observed. Results of the pilot survey indicated that traditional visual and acoustical survey methods for cetaceans should be adapted to find the elusive Baiji in the river. Currently, the range-wide Yangtze cetacean survey is in preparation. The survey will cover over 1,700 km of the Yangtze River from Yichang to Shanghai, and is expected to provide detailed information on Baiji and porpoise numbers and distribution patterns in the river. Discussion. Although the short-term goal of ex situ conservation is to rescue cetaceans from the river and to establish viable breeding populations in semi-natural reserves and in captivity, the long-term goal of releasing the animals back into the river when the threats have decreased and the natural environment has been improved, should not be neglected. Moreover, the in situ conservation efforts in the natural reserves, and even in the entire Yangtze River system, including the lakes, should not be ignored or abandoned at any time. The activities contributing to the conservation of the Baiji and the porpoise in the wild have the incidental effect of benefiting the entire Yangtze ecosystem and other rare threatened species. The dynamics of the groups of porpoises in semi-natural reserves should be monitored continually, in order to guide the establishment of breeding groups of Baiji in these semi-natural reserves in the near future. Conclusions. Under the existing severely degraded conditions of the Yangtze system, the sharply fall populations of Baiji and porpoises will not be suspended in the foreseeable future. Therefore, ex situ conservation should be emphasized, and the severely threatened Baiji in the river should be removed and translocated to semi-natural reserves for establishing viable breeding populations. The successful program of capturing, translocating and maintaining finless porpoises in the Shishou semi-natural reserve has demonstrated its adequacy as an ex situ environment for cetaceans. Following the successful pilot survey in the river, the immediate range-wide Yangtze cetacean survey is proposed and is in preparation. The range-wide survey is expected to ensure that any remaining Baiji can be found reliably and captured successfully after the survey. Recommendations and Perspectives. During the range-wide survey, not only the Baiji but also the porpoise as well as their habitats should be investigated based on visual and acoustical methods that adapted to the river and the animals. Meanwhile, the current risk levels to the Baiji and porpoises should be evaluated at each area where Baiji or porpoises can be reliably sighted. Any capture efforts should be targeted on the most threatened areas, or where there is maximum risk of injury or death. The immediate track of the Baiji should be carried out once a Baiji is sighted during the range-wide survey in order to obtain the movement route of the animals, which is crucial information for the successful capture operation. Additionally, the need to establish new semi-natural reserves for the porpoises should be placed on the agenda of local and central governments in the near future.

Aquatic resource conservation - The first Yangtze finless porpoise successfully born in captivity

Background. As the sole freshwater subspecies of finless porpoise (Neophocaena phocaenoides), the Yangtze finless porpoise (N. p. asiaeorientalis) lives only in the middle and lower reaches of the Yangtze River and its appended Poyang and Dongting Lakes. As a result of human activity on the river, including over and illegal fishing, pollution, transportation and dam construction, the population of Yangtze finless porpoises has been steadily and rapidly decreasing during the past several decades, which leads the animal to be endangered. Methods. For saving this unique animal from extinction, three corresponding measures, in situ conservation, ex situ conservation, and intensifying breeding research in captivity, were proposed and have been implemented since the 1980s. Results. After successfully rearing the animals in captivity for almost nine years, the first Yangtze finless porpoise was successfully born in captivity on July 5, 2005. The calf is male, with a body length of 69 cm. This is the first freshwater cetacean ever born in captivity. Conclusion. The successful birth of this calf confirms that it is possible to breed the Yangtze finless porpoise in captivity. Furthermore, this will greatly benefit the conservation efforts, and also greatly bolster our on-going efforts to study the reproductive biology of these animals. Recommendation. More studies and efforts are expected to establish a sustainable, captive colony of the Yangtze finless porpoise, which will not only greatly expand our knowledge about the reproduction biology of this animal, but also help to redeem the wild population through a careful yearly 'soft releasing' process.

Pollinator shift and reproductive performance of the qinghai-tibetan plateau endemic and endangered Swertia przewalskii (Gentianaceae)

Reproductive failure results in many plant species becoming endangered. However, little is known of how and to what extent pollinator shifts affect reproductive performance of endangered species as a result of the artificial introduction of alien insects. In this study we examined breeding systems, visitor species, visiting frequency and seed set coefficients of Swertia przewalskii in two years that had different dominant pollinator species (native vs. alien). Flowers of this species were protandrous and herkogamous and insects were needed for the production of seeds. The stigmatic receptivity of this species was shorter than for other gentians. No significant difference in seed set coefficient was found for hand-pollinated plants between the two years, indicating that pollinator shift only had a minor effect on this plant's breeding system. The commonest pollinators in 2002 were native bumblebees, alien honeybees and occasional solitary bees, however, only alien honeybees were observed in 2004. The flower visitation rate in both years was relatively high, although the total visit frequency decreased significantly in 2004. The control flowers without any treatment produced significantly fewer seed sets in 2004 than in 2002. In the past decade the seed production of this species may have partly decreased due to pollination by alien honeybees, however, we suggest that they might have acted as alternative pollinators ensuring seed production of S. przewalskii when native pollinators were unavailable. The main reason that this plant is endangered is probably the result of habitat destruction, but changes in land use, namely intensified agricultural practice and unfavorable animal husbandry have also contributed to its decline. We recommend that in-situ conservation, including the establishment of a protected area, is the best way to preserve this species effectively.

Simple procedures to assess and compare the ecological niche of species

Hutchinson's (1957; Cold Spring Harbour Symp Quant Biol 22:415-427) niche concept is being used increasingly in the context of global change, and is currently applied to many ecological issues including climate change, exotic species invasion and management of endangered species. For both the marine and terrestrial realms, there is a growing need to assess the breadth of the niches of individual species and to make comparisons among them to forecast the species' capabilities to adapt to global change. In this paper, we describe simple non-parametric multivariate procedures derived from a method originally used in climatology to (1) evaluate the breadth of the ecological niche of a species and (2) examine whether the niches are significantly separated. We first applied the statistical procedures to a simple fictive example of 3 species separated by 2 environmental factors in order to describe the technique. We then used it to quantify and compare the ecological niche of 2 key-structural marine zooplankton copepod species, Calanus finmarchicus and C. helgolandicus, in the northern part of the North Atlantic Ocean using 3 environmental factors. The test demonstrates that the niches of both species are significantly separated and that the coldwater species has a niche larger than that of its warmer-water congeneric species.

Fishery Exploitation and Stock Assessment of the Endangered Nassau Grouper, Epinephelus Striatus (Actinopterygii: Perciformes: Serranidae), in the Turks and Caicos Islands

The Nassau grouper, Epinephelus striatus (Bloch, 1792), is an endangered species that has been historically overexploited in numerous fisheries throughout its range in the Caribbean and tropical West Atlantic. Data relating fishery exploitation levels to stock abundance of the species are deficient, and protective regulations for the Nassau grouper are yet to be implemented in the Turks and Caicos Islands (TCI). The goal of this study was to conduct a stock assessment and evaluate the exploitation status of the Nassau grouper in the TCI. Materials and methods. Calibrated length cohort analysis was applied to published fisheries data on Nassau grouper landings in the TCI. The total lengths of Nassau groupers among the catches of spearfishers, lobster trappers, and deep sea fishers on the island of South Caicos during 2006 and 2008 were used with estimates of growth, natural mortality, and total annual landings to derive exploitation benchmarks. Results. The TCI stock experienced low to moderate fishing mortality (0.28, 0.18) and exploitation rates (0.49, 0.38) during the period of the study (2006, 2008). However, 21.2%-64.4% of all landings were reproductively immature. Spearfishing appeared to contribute most to fishing mortality relative to the use of lobster traps or hydraulic reels along bank drop-offs. Conclusion. In comparison with available fisheries data for the wider Caribbean, the results reveal the TCI as one of the remaining sites, in addition to the Bahamas, with a substantial Nassau grouper stock. In light of increasing development and tourism in the TCI, continued monitoring is essential to maintain sustainable harvesting practices.

High levels of genetic structuring as a result of population fragmentation in the tropical tree species Caesalpinia echinata Lam.

We have investigated levels of genetic diversity within and among seven remnant populations of Caesalpinia echinata Lam., an endangered species found as fragmented populations in three major areas around the coastal regions of Brazil. Using amplified fragment length polymorphism (AFLP) genetic markers, we detected levels of within-population genetic diversity ranging from 0.092 to 0.163, with the lowest values generally being found in the smallest populations. Estimates of between-population genetic differentiation were strongly correlated with geographical distance ( r = 0.884, p <0.001), which, along with a neighbour-joining phylogenetic analysis, strongly suggested high levels of genetic isolation by distance. Over half (62%) of the total genetic diversity was partitioned between populations, further highlighting the genetic distinctness of individual populations. Taken together, these results suggest that fragmentation has led to an increase in population differentiation between fragments of C. echinata. These formations will be of great value in the development of conservation plans for species exhibiting high levels of genetic differentiation due to fragmentation, such as indication of conservation unit size, which populations should be chosen as priority in conservation plans and which samples should be introduced in areas with a low number of individuals of brazilwood.

Indicators of the impact of climate change on migratory species

The Bonn Convention on the Conservation of Migratory Species of Wild Animals adopted a Resolution in 2005 recognising the impacts of climate change on migratory species. It called on Contracting Parties to undertake more research to improve our understanding of these impacts and to implement adaptation measures to reduce foreseeable adverse effects. Given the large diversity of taxa and species affected by climate change, it is impossible to monitor all species and effects thereof. However, it is likely that many of the key ecological and physical processes through which climate change may impact wildlife could be monitored using a suite of indicators, each comprising parameters of species/populations or groups of species as proxies for wider assemblages, habitats and ecosystems. Herein, we identify a suite of 17 indicators whose attributes could reveal negative impacts of climate change on the global status of migratory species: 4 for birds, 4 for marine mammals, 2 for sea turtles, 1 for fish, 3 for land mammals and 3 for bats. A few of these indicators would be relatively straightforward to develop, but most would require additional data collation, and in many cases methodological development. Choosing and developing indicators of the impacts of climate change on migratory species is a challenge, particularly with endangered species, which are subject to many other pressures. To identify and implement conservation measures for these species, indicators must account for the full ensemble of pressures, and link to a system of alerts and triggers for action.

Removing the threat of diclofenac to critically endangered Asian vultures

Veterinary use of the nonsteroidal anti-inflammatory (NSAID) drug diclofenac in South Asia has resulted in the collapse of populations of three vulture species of the genus Gyps to the most severe category of global extinction risk. Vultures are exposed to diclofenac when scavenging on livestock treated with the drug shortly before death. Diclofenac causes kidney damage, increased serum uric acid concentrations, visceral gout, and death. Concern about this issue led the Indian Government to announce its intention to ban the veterinary use of diclofenac by September 2005. Implementation of a ban is still in progress late in 2005, and to facilitate this we sought potential alternative NSAIDs by obtaining information from captive bird collections worldwide. We found that the NSAID meloxicam had been administered to 35 captive Gyps vultures with no apparent ill effects. We then undertook a phased programme of safety testing of meloxicam on the African white-backed vulture Gyps africanus, which we had previously established to be as susceptible to diclofenac poisoning as the endangered Asian Gyps vultures. We estimated the likely maximum level of exposure (MLE) of wild vultures and dosed birds by gavage (oral administration) with increasing quantities of the drug until the likely MLE was exceeded in a sample of 40 G. africanus. Subsequently, six G. africanus were fed tissues from cattle which had been treated with a higher than standard veterinary course of meloxicam prior to death. In the final phase, ten Asian vultures of two of the endangered species (Gyps bengalensis, Gyps indicus) were dosed with meloxicam by gavage; five of them at more than the likely MLE dosage. All meloxicam-treated birds survived all treatments, and none suffered any obvious clinical effects. Serum uric acid concentrations remained within the normal limits throughout, and were significantly lower than those from birds treated with diclofenac in other studies. We conclude that meloxicam is of low toxicity to Gyps vultures and that its use in place of diclofenac would reduce vulture mortality substantially in the Indian subcontinent. Meloxicam is already available for veterinary use in India.

Study of baleen whales’ ecology and interaction with maritime traffic activities to support management of a complex socio-ecological system

Thesis written in co-mentorship with Robert Michaud.

A Comparative study of germination strategies of two species of genus Allium sect. Allium

Germination experiments were performed with seeds of two species of genus Allium section Allium, a rare and endangered species A. pyrenaicum and a common A. sphaerocephalon. Different pre-treatments and a photoperiod of 24 h darkness were applied in order to simulate different germination conditions. Both species showed a high percentage of viable seeds a part of which were dormant. An elevate percentage of dormant seeds could be caused by a later collection time. Low altitude populations had more mortality than the others, possibly caused by the hard summer conditions during flowering and fruiting time. Comparisons between dates of species coexistence localities only show inter-population variability and it could be caused by the detected dormancy. Darkness accelerates germination, possibly for elongation radicle stimulation. Heat-shock pre-treatments decreased germination time in seeds from localities where fire is a probable event. The rarity of A. Pyrenaicum not seems to be caused by restricted germination requirements but is attributable to distinct habitat preferences, related to his altitudinal range of distribution