Edward H. Miller

Memorial University of Newfoundland · Department of Biology

We examined the ultrastructure of the mammalian os penis at the high‐resolution synchrotron level. Previously, bacular microanatomy had only been investigated histologically. We studied the baculum of the harp seal ( Pagophilus groenlandicus ), in which the baculum varies more in size and shape than does a mechanically constrained bone (humerus). W...

Growth, allometry, and characteristics of a sexually selected structure in wolverine (Gulo gulo (Linnaeus, 1758)), northern river otter (Lontra canadensis (Linnaeus, 1758)), and sea otter (Enhydra lutris (Linnaeus, 1758))

We studied calls of three sandpiper species to document species’ similarities and differences. We hypothesized that functionally different calls would differ in degree of divergence. We studied two nuptial calls (complex “Song” and rhythmically repeated aerial call, RRC) of males, and a general-purpose call (“alarm” trill, AT) given by parents of b...

Allometric analyses of sexually selected structures have revealed many patterns of evolutionary and behavioural significance, for example, in weapons, ornaments, and genitalia. We investigated allometry of the baculum (penis bone) relative to body size in post-growth adults of three large mustelids: wolverine (Gulo gulo (Linnaeus, 1758)), northern...

Acoustic communication is critical during early life phases in precocial birds, for example, adult alarm calls can elicit antipredator behaviour in young, and chick vocalisations can communicate information to parents about chick identity, condition, location, sex, or age. We opportunistically recorded Red‐capped Plover Charadrius ruficapillus and...

Birds often vocalize when threatened or captured by a predator. We present detailed qualitative analyses of calls from 24 red-capped plover (Charadrius ruficapillus) and 117 masked lapwing (Vanellus miles) chicks (Charadriidae) that we recorded during handling. Calls were structurally complex and differed between species. Calls showed moderate stru...

Acoustic communication is critical during early life phases in precocial birds. For example, adult alarm calls can elicit antipredator behaviour in young, and chick vocalisations can communicate information to parents about chick identity, condition, location, sex, or age. We investigated whether chick calls of two species of Australian Charadriida...

Behavioral ecology; communication; sexual dimorphism; social organization; walrus; weapon.

Communication; sexual dimorphism; sexual selection; social behavior; social organization; walrus; weapon

Large tusks characterize the extant walrus (Odobenus rosmarus) and its extinct relatives. Those socially selected organs vary between the walrus and fossil relatives, intraspecifically, and between the sexes. Tusks are used in innumerable interactions on land and in water, including fights during rut. “Play fighting” appears even in young calves th...

Natural and social selection are among the main shapers of biological diversity but their relative importance in divergence remains understudied. Additionally, although neutral evolutionary processes may promote phenotypic divergence, their potential contribution in speciation is often overlooked in studies of comparative morphology. In this study,...

La Becasina Común (Gallinago paraguaiae) posee dos subespecies, paraguaiae al este de los Andes a lo largo de Sudamérica y al norte de la Patagonia, y magellanica en centro y sur de Chile y en la Patagonia de Argentina Evaluamos la taxonomía en esta becasina analizando distintos despliegues acústicos utilizados durante la época reproductiva a lo la...

Acoustic displays of South American Gallinago p. paraguaiae and G. p. magellanica differ even more than between recognized sister species (e.g., G. gallinago, G. delicata). The 2 forms do not overlap geographically. We recommend that they be recognized as separate species.

Nuptial signals of related species can diverge rapidly, but this is not always the case. For example, homologous nuptial displays are recognizable even in shorebird lineages that have been separated for millions of years. We investigated similarities and differences in two nuptial and one non-nuptial (parental “alarm trill”, AT) sandpiper displays...

We analyzed breeding sounds of the two subspecies of South American Snipe Gallinago p. paraguaiae and G. p. magellanica to determine whether they may be different species: loud vocalizations given on the ground, and the tail‐generated Winnow given in aerial display. Sounds of the two taxa differ qualitatively and quantitatively. Both taxa utter two...

In mammals, males generally are larger than females, though such sexual-size differences have been documented primarily in adults and are relatively poorly known in early life. We studied sexual-size differences in pups of the northern elephant seal (Mirounga angustirostris (Gill, 1866)), which in adulthood is one of the most sexually dimorphic mam...

Most birds vocalize with an open beak, but vocalization with a closed beak into an inflating cavity occurs in territorial or courtship displays in disparate species throughout birds. Closed-mouth vocalizations generate resonance conditions that favor low-frequency sounds. By contrast, open-mouth vocalizations cover a wider frequency range. Here we...

We describe the display flight of the Andean Negrito (Lessonia oreas). The display flight consists of three parts: perched tsi notes, diagonal flight with tic notes, and turn and descent with trrrrrrrrrr wing-whirr and psie note. The wing-whirr appears to be a loud mechanical sound heretofore unknown in the species but consistent with the species'...

Least Sandpipers (Calidris minutila) were studied on the Queen Charlotte Islands, British Columbia, over five breeding seasons. Brood amalgamation and care of foreign chicks occur regularly, though at low incidence, among the approximately 90 breeding pairs there. Such behavior is probably important to chick survival and has little impact on adult...

Growth and size-scaling of the baculum and testes in the moderately polygynous hooded seal (Cystophora cristata) were studied using 107 specimens of known age (1 month to 28 years) from the northwestern Atlantic. Bacular growth was rapid between 2 and 5 years of age: length increased 150% and "density" (i.e., mass/length) increased 8-fold and mass...

Male walruses, Odobenus rosmarus (L.), summering (outside the breeding season) at an Alaskan hauling ground, use all available kinds of beach habitat: cobble and boulder beaches, rock benches, and large boulders. Formation and dissolution of large herds on land can occur rapidly. Walruses are very gregarious and positively thigmotactic. In cool wea...

Adult males of the grey seal, Halichoerus grypus (Fab.) (Phocidae), display the large snout conspicuously in threats. The male's snout has probably evolved through sexual selection as a visual display organ, as in some other extant phocids: the hooded seal, Cystophora cristata (Nilsson) and the two species of elephant seals, Mirounga (Gray). This m...

Display flights (.DF's) in the least sandpiper, Calidris minutilla (Vieill.), are described and their relations to ecological anti social variables assessed. DF activity is highest in early spring and in the morning, and is low in strong winds and poor visibility (fog). DF's are given only by males and generally cease upon pairing. Unpaired males m...

The complex aerial song of the Short-billed Dowitcher (Limnodromus griseus) is described, and recordings from northern British Columbia (subspecies caurinus), northern Manitoba (hendersoni), and Labrador (griseus) are analyzed. The song is a stereotyped sequence of one to five units that are repeated rapidly and increase successively in duration. E...

Sable Island, Nova Scotia, is the southernmost significant nesting area of the Least Sandpiper. Many birds nest around a single pond complex, which supports a lush vegetation that is heavily grazed by horses. Nests occur there and in nearby dry, sparsely vegetated habitat. Birds start arriving by mid-May (males first), and clutches (including repla...

Breeding Least Sandpipers were studied in Nova Scotia (1975–1976), Manitoba (1978), Yukon Territory (1979), and British Columbia (1982–1984). The most detailed study was on Sable Island, Nova Scotia, which is near the southern limit of the breeding range. The nesting period there was relatively long, and the species was strictly monogamous. Males t...

This chapter discusses territoriality in marine mammals. Territoriality refers to the exclusive use of fixed space, which entails obtaining, defending, or advertising occupancy of the space. Territoriality involves complex behavioral patterns used repeatedly in interaction and communication, which emerge in play early in life. The complex underwate...

We categorized and quantified the complete vocal repertoires of breeding adult auklets (Aethiini, 5 species) in their breeding areas to provide a baseline for comparative study of the structure and function of vocalizations within this monophyletic group of seabirds. We recognized 22 call types across species and 3-5 call types for each species. Ca...

We investigated molar-crown-size variation, sexual dimorphism, and allometry in the black bear (Ursus americanus), using hunter-shot specimens (n ¼ 429) from the island of Newfoundland, and museum specimens from elsewhere in Canada and the continental United States (n ¼ 502). We predicted higher variation in and weaker correlations among molar size...

The metabolic rate of harp (Pagophilus groenlandicus), harbor (Phoca vitulina), and ringed seals (Pusa hispida) was measured at various temperatures in air and water to estimate basal metabolic rates (BMRs) in these species. The basal rate and body composition of three harp seals were also measured throughout the year to examine the extent to which...

Species and sexual differences in vocalizations and the vocal tract are widespread. We studied the vocal tract of Common (Uria aalge) and Thick-billed (U. lomvia) Murres. We predicted anatomical or allometric differences in adults between species and sexes due to vocal differences related to social behavior (Common Murres nest at higher densities;...

Previous studies have suggested that diets of river otters (Lontra canadensis) vary in response to seasonal shifts in prey availability, and that they select slowly moving fish of moderate size. To test these assumptions for marine-coastal river otters in Newfoundland, Canada, we reconstructed diets and estimated body length of important fish prey...

Many sexually-selected structures are variable and positively allometric relative to body size. For the western martenMartes caurina Merriam, 1890 from Vancouver Island, British Columbia, we investigated variation in the baculum compared with other bones and allometry of bacular to body size. Carcass length did not differ, and humeral and mandibula...

Vocalizations are an important component of male elephant seal agonistic behaviour. Acoustic and behavioural components of vocalizations emitted during agonistic contests show gross differences between young and old males, but the variation with age depends on the specific feature. Vocalizations become more frequent and effective at later ages. Aco...

Pre- and post-weaning functional demands on body size and shape of mammals are often in conflict, especially in species where weaning involves a change of habitat. Compared with long lactations, brief lactations are expected to be associated with fast rates of development and attainment of adult traits. We describe allometry and growth for several...

Intraspecific sexual differences, high variation, and positive allometry of sexually-selected external display structures are common. Many sexually-selected anatomical specializations occur in the avian vocal tract but intraspecific variation and allometry have been investigated little. The tracheal bulla bulla syringealis occurs in males of most d...

Pinnipeds generally swallow prey whole, and most have simple, homodont, nonoccluding cheek teeth. We investigated whether cheek teeth in seals are more variable and weakly integrated than in terrestrial Carnivora. We measured mandibular length and crown length of mandibular postcanines (PCs) in ringed seals (Pusa hispida; n ¼ 912) from the Canadian...

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10months, AFS: 4 months), even under local sympat...

In this first study of vocal individuality in the Charadriidae, we describe vocal variation in the endangered Piping Plover (Charadrius melodus). We studied a long-distance advertisement call given by breeding males during aerial displays, because functionally similar vocalizations of scolopacids are known to be individualistic. We analyzed recordi...

In this first study of vocal individuality in the Charadriidae, we describe vocal variation in the endangered Piping Plover (Charadrius melodus). We studied a long-distance advertisement call given by breeding males during aerial displays, because functionally similar vocalizations of scolopacids are known to be individualistic. We analyzed recordi...

In the most common models of communication, it is assumed that animals provide reliable information about phenotype, and hence can settle competitive contests without physical interactions like fights. This assumption has rarely been tested for wild mammals. Recent studies of mammals have revealed relationships of vocal attributes to age and body s...

Vocal learning in mammals is sparsely documented, and there are few reports of vocal learning by wild mammals. In particular, no information based on longitudinal data for identified individuals exists, even for well-studied highly social species in which vocal communication is an important aspect of social life. We present such information for the...

The proboscis of male elephant seals (Mirounga Gray, 1827) has been suggested as an example of a secondary sexual trait since Darwin. There has been much speculation about its social function (e.g., optical signal of breeding status, amplification of vocalizations, cue for female choice). However, it has never been studied in detail, probably becau...

We studied breeding vocalizations and the vocal repertoire of the endangered piping plover (Charadrius melodus Ord, 1824) to describe diversity of the species' non-learned vocalizations, provide a basis for comparative studies, and enable standardization of terminology and interpretation of vocal classes for management purposes. Adults have 14 call...

Structure and growth of the baculum (os penis) in arctoid carnivores have been well described for many species. This study presents the first extensive analysis of bacular growth and variation for bears (Ursidae), based on 871 bacula of polar bears Ursus maritimus (858 of known age) that were shot in the Canadian Arctic from 1994 to 1997. Bacular l...

The biology of the endangered Tuamotu Sandpiper (Prosobonia cancellata) is essentially unknown. We analyzed vocalizations from presumed adult individuals and pairs, and family groups, recorded in French Polynesia during March 1990 and 2003. We recognized three types of vocalizations. Presumed adults uttered types I and II. These were brief (about 3...

Predation on eggs, nestlings, and breeding adults of Red-naped Sapsuckers, Sphyrapicus nuchalis, Northern Flickers, Colaptes auratus, Hairy Woodpeckers, Picoides villosus, and Williamson's Sapsuckers, S. thyroideus, was documented in the Hat Creek valley, south-central British Columbia from 1989-1994. Predation by Black Bears (Ursus americanus), De...

We compared allometry and variation in the baculum (os penis), mandible, and humerus of the harp seal, Pagophilus groenlandicus. This species is presumed to have a promiscuous mating system in which choice of mate by females during intromission with different males is likely. The baculum is large and grows throughout life so may be an honest indica...

We compared allometry and variation in the baculum (os penis), mandible, and humerus of the harp seal, Pagophilus groenlandicus. This species is presumed to have a promiscuous mating system in which choice of mate by females during intromission with different males is likely. The baculum is large and grows throughout life so may be an honest indica...

Bacula are relatively small in terrestrially mating species of pinnipeds (otariids and elephant seals, Mirounga), perhaps reflecting adaptive size reduction to minimize bacular fracture. Fur seals and sea lions (Otariidae) are a good group with which to investigate this question, because most species copulate solely on land and body size varies int...

Bacula rue relatively small in terrestrially mating species of pinnipeds (otariids and elephant seals, Mirounga), perhaps reflecting adaptive size reduction to minimize bacular fracture. Fur seals and sea lions (Otariidae) are a good group with which to investigate this question, because most species copulate solely on land and body size varies int...

The extent and ecological significance of time spent (“hauled out”) on ice by phocid seals are poorly known. In this study, haulout of nine captive harp seals (Pagophilus groenlandicus) was monitored and quantified with recorders attached to the seals. The recorders monitored haulout (presence in or out of salt water). Data were collected continuou...

A bstract The baculum in Arctocephalus p. pusillus reaches up to 14.1 cm in length, 13.5 g in mass, and 1.3 g/cm in density (= mass/length). A pubertal growth spurt occurs between 2 and 3 yr of age, when bacular length increases by 28%, mass by 124%, and density by 77%; concurrently, body length increases by 14%. A second, weaker spurt occurs at so...

Growth and size-scaling of the baculum and testes in the moderately porygynous hooded seal (Cystophora cristata) were studied using 107 specimens of known age (1 month to 28 years) from the northwestern Atlantic. Bacular growth was rapid between 2 and 5 years of age: length increased 150% and "density" (i.e., mass/length) increased 8-fold and mass...

Plumage, size, and sexual dimorphism of the endemic Hairy Woodpecker subspecies Picoides villosus picoideus are described and compared with a less isolated insular population. Museum specimens of P. v. picoideus from the Queen Charlotte Islands, British Columbia were compared with P. v. harrisi specimens from Vancouver island, British Columbia. Spe...

We investigated quantitative relationships of bacular size to age, length of body, and testicular size in the harp seal (Pagophilus groenlandicus), based on collections made in the northwestern Atlantic from 1985 to 1992. Bacular growth was faster than growth in length of body except in old seals (>9 years of age), in which relative growth was isom...

Prey energy density values are crucial inputs to bioenergetic consumption models. Vertebrate predators in the northwest Atlantic consume a variety of prey species, but the proximate composition (PC; proportions of lipid, protein, ash and water) and energy density (ED; kJ g(-1)) of prey, and their variability, are known poorly. In this study, key pr...

Digestive efficiency (DE) is influenced by many factors including food type or quality. Assimilation efficiency (AE) and DE of 12 captive harp seals (Phoca groenlandica) were estimated for five prey types in large outdoor seawater tanks. In trials of >9 days' duration, the seals were fed Atlantic cod (Gadus morhua), Arctic cod (Boreogadus saida), G...

Faeces were collected from four captive harbour seals (Phoca vitulina) that consumed known amounts of herring (&pea harengus), walleye pollock (Therugru chalcogrumma), Pacific hake (Merluccius productus), surf smelt (Hypomesus pretiosus), and juvenile chinook salmon (Oncorhynchus tshmvytschu). The goal was to determine which structures (hard parts)...

Acoustic communication is important in the social life of pinnipeds, but our knowledge of it is limited. Physical analyses of sound signals are unavailable for some species, and are scant for others (Watkins and Wartzok, 1985; Table 1). Most existing analyses cover only a small portion of each species’ repertoire, are not very detailed, and provide...

The Black Oystercatcher (Haematopus bachmani) is a year-round resident with an extensive range from southern California to Alaska. Because of the size of this range (about 28° of latitude) and its simple, nearly linear shape, we hypothesized that the species would exhibit adaptive geographic variation in nesting phenology and clutch size. Museum eg...

Communication is complex and variable in long-lived and adaptable species. In such species, communication is shaped by learning, personal histories, variable social and ecological circumstances, and bonds between individuals. It is also often extremely subtle, especially in highly social species, in which individuals continually appraise and respon...

Vocalizations of Baird's Sandpiper were recorded in northeastern Alaska, U.S.A. Males beat the wings continuously in a loose deep flutter during display flights. They utter Rhythmically Repeated Calls (RRCs), Song and Chatter in display flights, and a Laugh when mildly disturbed. RRCs averaged 690 ms long with intervals between them of 280 ms. They...

Ethology arose as a discipline within zoology, with distinctly different aims and methods from psychology (Tinbergen, 1963). From its inception, ethology embraced phylogenesis, inheritance, “survival value,” and adaptation—research areas that relied on comparative observations and inductive reasoning (Darwin, 1859, 1872). Our understanding of anima...

Breeding St&birds (Aphriza virgata) were studied in the Yukon Territory and Alaska. They utter three common call types: Song, Rhythmically Repeated Call (RRC), and Laugh. RRCs are emitted as bouts during wide-ranging aerial displays, presumably by males. Each RRC is composed of a few brief pulses followed by a long, nearly constant-frequency portio...

Unpaired male least sandpipers (Calidris minutilla) emitted three main kinds of calls to attract mates and in sexual or agonistic interactions. The call types varied slightly across the nesting range (samples from British Columbia, Yukon Territory, Manitoba, and Nova Scotia). One call type, analyzed in most detail, was given in long rhythmic series...

Walruses spend much time hauled out on land or ice where they engage in numerous short-range social interactions within densely packed groups. Almost all interactions include sounds. Airborne sounds include approx 10 structurally defined classes, 6 of which occur commonly: 3 are used in threat, 2 occur between females and their offspring, and one (...

Shorebirds offer rich opportunities for comparative research because of their wide geographic distribution and their phylogenetic, ecological, and social diversity. There are more than 200 extant species, which occur in all zoogeographic regions of the world. Some species are resident and sedentary year-round, like the Shore Plover (Charadrius nova...

Display Flights (DFs) of the Least Sandpiper (Calidris minutilla) are described. DFs were prolonged flights (averaging 1.2 min in Manitoba and 3.4 min in Nova Scotia), about 10-30 m in altitude, by unmated males over their future nesting areas. Displaying males alternated brief glides and bouts of rapid shallow wingbeats (Flutters) throughout DFs;...

Sandpipers in the subfamily Calidridinae (Scolopacidae) breed in the Arctic and Subarctic. In all species, unpaired males engage in distinctive aerial displays. Such displays are described for three species: Dunlin (Calidris alpina L.), Semipalmated Sandpiper (Calidris pusilla L.), and Stilt Sandpiper (Calidris himantopus Bonaparte). In each of the...

The purpose of this paper is to explore some techniques of representing and analyzing sound spectrograms, with reference to detecting and characterizing graded vocalizations. Description and classification of sounds based on their structure can be separate from research into their functions, and this paper considers affinities among calls as determ...

Social behavior of male Pacific walruses. Odobenus rosmarus (L.), summering (i.e. outside the breeding season) on an Alaskan hauling ground is described. Social interaction on land is mostly agonistic. Visual presentation of tusks and striking with tusks feature prominently in most agonistic interactions: vocal communication occurs in a minority of...

Fur seals Arctocephalus forsteri and walruses Odobenus rosmarus show similar facial expressions in a variety of social and non-social contexts. In non-social settings, both species modify the facial appearance by erecting the mystacial vibrissae while grooming the forequarters, while yawning, and during olfactory/tactile investigation of objects. D...