OBSERVATIONS OF FLORIDA MANATEES(TRICHECHUS MANATUS LATIROSTRIS) AROUND SELECTED POWER PLANTS IN WINTER

Sixty-four aerial surveys were flown during cold, winter weather between December 1982 and February 1772 to provide an index of trends in abundance of the Florida manatee ( Trichechus manatus latirostris )around five Florida Power & Light Company (FPL) plants. The surveys documented high winter use of warm-water discharges at four of the plants (Canaveral, Riviera. Port Everglades, and Fort Myers). The total number of manatees sighted at the five primary plants surveyed during a single survey ranged from 6 to 765 manatees (= 330 ± 45 manatees per survey): thus, up to 41% of the total Florida manatee minimum population of 1.850 animals may be found on cold days at FPL plants. The number of calves sighted at the plants during winter has decreased; similarly, for three consecutive years (winter 1989–1970 through winter 1991–1992), the percentage of the manatees that are calves sighted at the plants has decreased. Although data from the surveys of FPL plants do not necessarily reflect what may be happening with manatees located elsewhere in Florida, a possible reduction in calf numbers, coupled with increasing numbers of dead calves being documented by scientists involved in manatee carcass salvage work, is cause for concern in this slow-reproducing, endangered species.     

DETERMINATION OF MANATEE POPULATION TRENDS ALONG THE ATLANTIC COAST OF FLORIDA USING A BAYESIAN APPROACH WITH TEMPERATURE-ADJUSTED AERIAL SURVEY DATA

In many animal population survey studies, the construction of a stochastic model provides an effective way to capture underlying biological characteristics that contribute to the overall variation in the data. In this paper we develop a stochastic model to assess the population trend and abundance of the Florida manatee, Trichechus manatus latirostris , along the Atlantic coast of the state, using aerial survey data collected at winter aggregation sites between 1982 and 2001. This model accounts for the method by which the manatees were counted, their movements between surveys, and the behavior of the total population over time. The data suggest an overall increase in the population from 1982 to 1989 of around 5%–7%, a reduction in growth or a leveling off (0%–4% annual growth) from 1990 to 1993, and then an increase again of around 3%–6% since 1994. In winter 2001–2002 (the most recent survey season for which analyses were done), we estimated the adult manatee population along the east coast of Florida to be 1,607 individuals (range = 1,353–1,972; 95% credible interval). Our estimate of manatee abundance corresponds well with maximum counts (approximately 1,600 manatees) produced during synoptic aerial surveys under optimal conditions. Our calculations of trends correspond well with mark and recapture analyses of trends in survival of adult manatees along the east coast through the early 1990s. Our population trend estimates since that time are more optimistic than those generated by mark-recapture models.     

AERIAL VIDEOGRAMMETRY FROM A TETHERED AIRSHIP TO ASSESS MANATEE LIFE-STAGE STRUCTURE

Aerial videogrammetry from an airship tethered to a boat was used to assess the life-stage structure of manatees in the Blue Waters area of the Homosassa River on the west coast of central Florida. Individual frames of videos were loaded onto a computer and lengths of manatees measured using graphics software. All images of manatees were measured independently by three observers. Measurements were transformed to total manatee lengths by calculating the ratio between the measurements of a reference object of known length and the measurement of the manatee. Length estimates were highly variable but were unbiased with respect to observer. Results of an analysis of variance suggested that the hypothesis that lengths differed among days and among observers should not be rejected. Life-stage structures were represented in three schemes. One scheme-which included four life stages: dependent calves, juveniles, animals in tramsition between juvenile and adult, and adults-was included to provide the most information without sacrificing confidence in the life-stage distribution. Overall, the video system was effective at capturing images of manatees for life-stage characterization and, with improvements in image resolution, could become a valuable tool for photo-identification in sight-resight experiments.     

PERMANENT GENETIC RESOURCES: Eighteen new polymorphic microsatellite markers for the endangered Florida manatee, Trichechus manatus latirostris

Here we describe 18 polymorphic microsatellite loci for Trichechus manatus latirostris (Florida manatee), isolated using a polymerase chain reaction‐based technique. The number of alleles at each locus ranged from two to four (mean = 2.5) in specimens from southwest (n = 58) and northeast (n = 58) Florida. Expected and observed heterozygosities ranged from 0.11 to 0.67 (mean = 0.35) and from 0.02 to 0.78 (mean = 0.34), respectively. Departures from Hardy–Weinberg equilibrium occurred at two loci. There was no evidence of genotypic disequilibrium for any pair of loci. For individual identification, mean random‐mating and θ‐corrected match probabilities were 9.36 × 10^?7 and 1.95 × 10^?6, respectively.     

REPRODUCTION IN FEMALE MANATEES OBSERVED IN SARASOTA BAY, FLORIDA

Resightings of recognizable individual Florida manatees, Trichechus manatus latirostris , in Sarasota Bay provided information on life history and reproductive parameters. These parameters have been documented at winter aggregation sites, but few data existed for Florida's west coast at non-winter sites. Life history data are needed to estimate annual reproduction and survival rates and construct population models for manatees along west central Florida. Observations of 113 recognizable manatees of known sex sighted between 1993 and 1997 indicated a 1: 1 sex ratio. Fifty-three females produced at least 55 calves during five years of observation. Two large, presumably sexually mature females were never observed with calves in over five years of monitoring. Not surprisingly, several reproductive parameters matched those measured in previous studies at winter sites with a mean of 1.8 seasons of calf dependence, 2.2 seasons between births, and age of sexual maturity at 3–4 yr. Birth rates, fecundity, and proportions of females with calves varied only slightly among investigations.     

Functional morphology of the taste buds of Florida manatee,Trichechus manatus latirostris

The Florida manatee, Trichechus manatus latirostris, is a fully aquatic, threatened marine mammal for which increased understanding of their physiology, reproduction, and nutrition supports management decisions. Manatees may use taste to distinguish saltwater gradients, toxin detection, food assessment, and social interactions. This study sought to locate and characterize manatee taste buds comparing location, structure, number, and size to other species. Entire heads from manatees (6 males, 4 females) of various ages were obtained. The muzzle, tissue surrounding the nares, oral cavity, and epiglottis were examined grossly for pits and papillae. Tissues were examined using light and transmission electron microscopy. Within the predominant taste bud location, the tongue root, taste bud number was estimated using samples from four animals. The average number of taste buds within the tongue root was 11,534 (range 2,711–23,237) with sparse taste buds located on the soft palate and epiglottis. The location along the lateral surface of the tongue root and bordered by grooves, through which tastants could be easily transported, has functional significance. Large numbers of taste buds within the tongue root suggest that taste is an important component of manatee sensory systems and behavioral research would clarify this.     

SURVIVAL ESTIMATES FOR FLORIDA MANATEES FROM THE PHOTO-IDENTIFICATION OF INDIVIDUALS

We estimated adult survival probabilities for the endangered Florida manatee ( Trichechus manatus latirostris ) in four regional populations using photoidentification data and open-population capture-recapture statistical models. The mean annual adult survival probability over the most recent 10-yr period of available estimates was as follows: Northwest - 0.956 (SE 0.007), Upper St. Johns River - 0.960 (0.011), Atlantic Coast - 0.937 (0.008), and Southwest - 0.908 (0.019). Estimates of temporal variance independent of sampling error, calculated from the survival estimates, indicated constant survival in the Upper St. Johns River, true temporal variability in the Northwest and Atlantic Coast, and large sampling variability obscuring estimates for the Southwest. Calf and subadult survival probabilities were estimated for the Upper St. Johns River from the only available data for known-aged individuals: 0.810 (95% CI 0.727–0.873) for 1st year calves, 0.915 (0.827–0.960) for 2nd year calves, and 0.969 (0.946–0.982) for manatee 3 yr or older. These estimates of survival probabilities and temporal variance, in conjunction with estimates of reproduction probabilities from photoidentification data can be used to model manatee population dynamics, estimate population growth rates, and provide an integrated measure of regional status.     

FORENSIC METHODS FOR CHARACTERIZING WATERCRAFT FROM WATERCRAFT-INDUCED WOUNDS ON THE FLORIDA MANATEE (TRICHECHUS MANATUS LATIROSTRIS)

Watercraft-related mortality represents 1,253 (24.9%) of 5,033 Florida manatee ( Trichechus manatus latirostris ) deaths recorded between 1 January 1979 and 31 December 2004. Wound patterns generated by collisions with watercraft are diagnostic. Sets of cuts and scrapes that are roughly equidistant and perpendicular to the direction of vessel travel are consistent with lacerations made by propeller blades. From these lesions, estimates of propeller diameter, pitch, rotation, and direction of travel may be obtained. Considerable overlap of propeller sizes and pitches on different size vessels, common use of counter rotation propellers, and numerous other complicating factors may confound efforts to accurately predict vessel size and type from propeller wounds. Of the more than one million watercraft registered in Florida, 98% are ≤12.2 m (40 ft), yet watercraft 5.3–36.6 m (17.5–120 ft) are known to have killed manatees. Analysis of a 5-yr subset of mortality data suggests that a disproportionate number of propeller-caused watercraft-related mortalities could be attributed to propeller diameters ≥43.2 cm (17 in.), inferring that these were caused by watercraft ≥12.2 m (40 ft).     

HABITAT SELECTION BY MANATEES IN SARASOTA BAY, FLORIDA

Habitat selection by Florida manatees ( Trichechus manatus latirostris ) is influenced by, among other things, availability of food, thermal, and freshwater resources. However, habitat selection by females with dependent calves may differ from that of other demographic groups with regard to the relative importance of these factors. Additional factors that may influence habitat selection for females with dependent calves include ambient noise, strong currents, or increased foraging requirements. We examined distributional data for manatees from aerial surveys of the coastal waters near Sarasota, Florida, between 1994 and 2004 to determine whether habitat selection by groups of manatees that included calves differed from that of other groups. We characterized groups according to their location within seven habitat types. Enclosed bays not traversed by the Intracoastal Waterway had the highest proportions of groups with calves. Groups with calves used a No Entry refuge (from which almost all human use is barred) to a greater extent than did other groups. Overall, groups with calves exhibited significantly different habitat selection from groups without calves ( P < 0.001, χ²= 43.0, df = 6), but this was not consistent across seasons. During the winter and spring, thermal requirements influenced manatees to such an extent that all demographic groups selected habitat similarly.     

SIMULATED VESSEL APPROACHES ELICIT DIFFERENTIAL RESPONSES FROM MANATEES

One of the most pressing concerns associated with conservation of the endangered Florida manatee is mortality and serious injury due to collisions with watercraft. Watercraft collisions are the leading identified cause of manatee mortality, averaging 25% and reaching 31% of deaths each year. The successful establishment and management of protected areas depend upon the acquisition of data assessing how manatees use different habitats, and identification of environmental characteristics influencing manatee behavior and habitat selection. Acoustic playback experiments were conducted to assess the behavioral responses of manatees to watercraft approaches. Playback stimuli made from prerecorded watercraft approaches were constructed to simulate a vessel approach to approximately 10 m in sea grass habitats. Stimulus categories were (1) silent control, (2) approach with outboard at idle speed, (3) vessel approach at planning speed, and (4) fast personal watercraft approach. Analyses of swim speed, changes in behavioral state, and respiration rate indicate that the animals responded differentially to the playback categories. The most pronounced responses, relative to the controls, were elicited by personal watercraft. Quantitative documentation of response during playbacks provides data that may be used as the basis for future models to predict the impact of specific human activities on manatees and other marine mammal populations.     

Characterization of highly informative cross-species microsatellite panels for the Australian dugong (Dugong dugon) and Florida manatee (Trichechus manatus latirostris) including five novel primers

The Australian dugong (Dugong dugon) and Florida manatee (Trichechus manatus latirostris) are threatened species of aquatic mammals in the order Sirenia. Sirenian conservation and management actions would benefit from a more complete understanding of genetic diversity and population structure. Generally, species-specific microsatellite markers are employed in conservation genetic studies; however, robust markers can be difficult and costly to isolate. To increase the number of available markers, dugong and manatee microsatellite primers were evaluated for cross-species amplification. Furthermore, one manatee and four dugong novel primers are reported. After polymerase chain reaction optimization, 23 (92%) manatee primers successfully amplified dugong DNA, of which 11 (48%) were polymorphic. Of the 32 dugong primers tested, 27 (84%) yielded product in the manatee, of which 17 (63%) were polymorphic. Dugong and manatee primers were compared and the most informative markers were selected to create robust and informative marker-panels for each species. These cross-species microsatellite marker-panels can be employed to assess other sirenian populations and can provide beneficial information for the protection and management of these unique mammals.     

DISTRIBUTION AND INNERVATION OF FACIAL BRISTLES AND HAIRS IN THE FLORIDA MANATEE (TRICHECHUS MANATUS LATIROSTRIS)

The distribution and anatomy of sirenian perioral bristles (modified vibrissae) and facial hairs are of interest because of their use during feeding and tactile exploration. In the present study we have identified six fields of perioral bristles on the face of the Florida manatee (T. manatus latirostris), four (U1-U4) on each side of the upper lips and oral cavity, and two (L1-L2) on each side of the lower lip pad, inside the oral cavity and rostral to the horny mandibular pad. Each field has a characteristic location, number of bristles, and range of bristle length and diameter. There is a mean of 110 (± 19) bristles per side, with no left-right differences. Branches of the infraorbital nerve innervate the bases of the largest bristles (U2 group) on the upper bristle pad, and the inferior alveolar nerve supplies the bristles of the lower bristle pad. The dorsal and ventral buccal branches of the facial nerve innervate the superficial facial musculature, which is likely to be involved in bristle eversion and other movements which constitute feeding behavior. Hair is denser in the facial region than on the remainder of the body. Within the face, hair is denser on the oral disk than on the supradisk. The oral disk contains bristle-like hair, whereas the supradisk region possesses hair that is similar in length and diameter to that on the postcranial body. The mean total of bristles and hairs per face was 1,942. Means for the subregions were 220 (± 39) bristles on the perioral bristle pads, 601 (± 115) bristlelike hairs in the oral disk region, 710 (± 229) typical hairs in the supradisk region, and 411 (± 108) typical hairs on the chin. There were no significant differences between left and right side counts. Facial hair density was inversely correlated with facial area and body size. These data provide new information on the anatomical basis of the exceptional orofacial activities characteristic of manatees during feeding and tactile exploration.     

Behavioral response of manatees to variations in environmental sound levels

Florida manatees (Trichechus manatus latirostris) inhabit coastal regions because they feed on the aquatic vegetation that grows in shallow waters, which are the same areas where human activities are greatest. Noise produced from anthropogenic and natural sources has the potential to affect these animals by eliciting responses ranging from mild behavioral changes to extreme aversion. Sound levels were calculated from recordings made throughout behavioral observation periods. An information theoretic approach was used to investigate the relationship between behavior patterns and sound level. Results indicated that elevated sound levels affect manatee activity and are a function of behavioral state. The proportion of time manatees spent feeding and milling changed in response to sound level. When ambient sound levels were highest, more time was spent in the directed, goal‐oriented behavior of feeding, whereas less time was spent engaged in undirected behavior such as milling. This work illustrates how shifts in activity of individual manatees may be useful parameters for identifying impacts of noise on manatees and might inform population level effects.     

PREHENSILE USE OF PERIORAL BRISTLES DURING FEEDING AND ASSOCIATED BEHAVIORS OF THE FLORIDA MANATEE (TRICHECHUS MANATUS LATIROSTRIS)

The use of perioral bristles (modified vibrissae) by 17 captive Florida manatees and approximately 20 wild manatees was analyzed. Captive manatees were fed six species of aquatic vegetation normally eaten in the wild (four freshwater species and two seagrasses). Inanimate objects were placed in the holding tanks with manatees at Lowry Park Zoological Gardens (Tampa, FL) to determine the degree to which perioral bristles were used in exploration and to define the range of manipulative behavior. In addition, behavioral observations were made on the use of perioral bristles during social interactions with conspecifics. Observations were recorded using a Hi8-format video camera. Florida manatees possess an unusually large degree of fine motor control of the snout and perioral bristles. The large and robust perioral bristle fields of the upper lip were used in a prehensile manner during feeding. Bristle use by manatees feeding on submerged vegetation differed from that seen during feeding on floating vegetation. Other behavioral use of the perioral bristles shows variation depending upon the situation encountered. The degree of plasticity of perioral bristle use supports our hypothesis that the vibrissal-muscular complex of the Florida manatee has evolved to increase the efficiency of grazing and browsing on aquatic vegetation and to fully maximize the potential of the manatee as a generalist feeder. The manipulative and sensitive nature of the manatee snout is likely a manifestation of a complex sensory and motor system which has evolved for marine mammal aquatic herbivores living in shallow turbid habitats.     

THE EVOLUTION OF SPERM SIZE IN BIRDS

Sperm size varies enormously among species, but the reasons for this variation remain obscure. Since it has been suggested that swimming velocity increases with sperm length, earlier studies proposed longer (and therefore faster) sperm are advantageous under conditions of intense sperm competition. Nonetheless, previous work has been equivocal, perhaps because the intensity of sperm competition was measured indirectly. DNA profiling now provides a more direct measure of the number of offspring sired by extrapair males, and thus a more direct method of assessing the potential for sperm competition. Using a sample of 21 species of passerine birds for which DNA profiling data were available, we found a positive relation between sperm length and the degree of extrapair paternity. A path analysis, however, revealed that this relationship arises only indirectly through the positive relationship between the rate of extrapair paternity and length of sperm storage tubules (SSTs) in the female. As sperm length is correlated positively with SST length, an increase in the intensity of sperm competition leads to an increase in sperm length only through its effect on SST length. Why females vary SST length with the intensity of sperm competition is not clear, but one possibility is that it increases female control over how sperm are used in fertilization. Males, in turn, may respond on an evolutionary time scale to changes in SST size by increasing sperm length to prevent displacement from rival sperm. Previous theoretical analyses predicting that sperm size should decrease as sperm competition becomes more intense were not supported by our findings. We suggest that future models of sperm-size evolution consider not only the role of sperm competition, but also how female control and manipulation of ejaculates after insemination selects for different sperm morphologies.     

AN EPIZOOTIC OF FLORIDA MANATEES ASSOCIATED WITH A DINOFLAGELLATE BLOOM

Over a 10-wk period in early 1982, 39 Florida manatees ( Trichechus manatus latirostris ) were found dead in the lower Caloosahatchee River and nearby waters of southwestern Florida. Two were killed by boats. The remainder showed no evidence of trauma. Lesions indicative of infectious agents were not identified, and bacteriological and contaminant residue findings were unremarkable. Nonspecific lesions of congestion and hemorrhage were identified in brain tissue. Numerous reports were also received of manatee morbidity. Some distressed manatees showed no biochemical lesions in clinical analyses of blood samples and recovered quickly. Timing of manatee illnesses coincided with fish and double-crested cormorant ( Phalacrocorax auritus ) mortality and morbidity. A widespread bloom of the dinoflagellate red tide organism ( Gymnodinium breve ) also coincided with these incidents. G. breve produces potent neurotoxins (brevetoxins). Circumstantial evidence links these events, and possible routes of exposure may include ingestion of filter-feeding ascidians. Ecological conditions that magnified the extent of the epizootic included an early dispersal of manatees into the area from a nearby winter aggregation site and unusually high salinities that facilitated the inshore spread of the red tide bloom. Management responses to future episodes of red tide in manatee areas are suggested.     

INDIVIDUAL VARIATION IN SPERM COMPETITION SUCCESS OF YELLOW DUNG FLIES,SCATOPHAGA STERCORARIA

Intraspecific variation in the proportion of offspring sired by the second male to mate with a female (P₂) is an aspect of sperm competition that has received little attention. We examined variation in the sperm competition success of individual male dung flies, Scatophaga stercoraria. In unmanipulated matings, copula duration was dependent on male size with smaller males copulating for longer. A principal component analysis was used to generate uncorrelated scores based on a male's size and copula duration. Using these scores demonstrated that P₂ values were dependent both on the relative size and copula durations of competing males. When copula duration was held constant, the success of an individual male increased as his body size, relative to the first male, increased. We interrupted copulations of “large” and “small” second males and fitted the resultant P₂ values to a linear model of sperm competition with unequal ejaculates. The data fit well to a model of sperm displacement in which sperm mix quickly on introduction to the sperm stores. Furthermore, they show that “large” males have a greater rate of sperm displacement than “small” males. The levels of prey availability during testis maturation may influence a male's success in sperm competition although his immediate mating history does not. We show why an understanding of variation in sperm competition success is important for understanding the mechanisms and evolutionary significance of sperm competition.     

INFERENCE OF SPERM COMPETITION FROM BROODS OF FIELD-CAUGHT DROSOPHILA

In field studies of multiple mating and sperm competition there typically is no experimental control over the number of times that a female mates, the interval between matings, or the genetic identity of multiple fathers contributing to a brood. Irrespective of this complexity, high-resolution molecular markers can be used to assign paternity with considerable confidence. This study employed two highly heterozygous microsatellite loci to assess multiple paternity and sperm displacement in a sample of broods taken from a natural population of Drosophila melanogaster. The large number of alleles present at each of the loci makes it difficult to derive explicit maximum-likelihood estimates for multiple paternity and sperm displacement from brood samples. Monte Carlo simulations were used to estimate maximum-likelihood parameters for the distribution of female remating frequency and the proportion of offspring sired by the second or subsequent mating males. Estimates were made based on genotypes scored at two distinct marker loci because they were found to give statistically homogeneous results. Fitting a Poisson distribution of number of matings, the mean number of males mated by a female was 1.82. The sperm displacement parameter estimated from doubly mated females were 0.79 and 0.86 for the two loci (0.83 for the joint estimate). The overall probability that a multiply mated female will be misclassified as singly mated was only 0.006, which indicates that microsatellites can provide excellent resolution for identifying multiple mating. In addition, microsatellites can be used to generate relatively precise estimates of sperm precedence in brood-structured samples from a natural population.     

SPERM COMPETITION SELECTS BEYOND RELATIVE TESTES SIZE IN BIRDS

Sperm morphology varies considerably across taxa, and postcopulatory sexual selection is thought to be one of the main forces responsible for this diversity. Several studies have investigated the effects of the variation in sperm design on sperm function, but the consequences of variation in sperm design on testis morphology have been overlooked. Testes size or architecture may determine the size of the sperm they produce, and selection for longer sperm may require concomitant adaptations in the testes. Relative testes size differs greatly between species and is often used as an index of sperm competition, but little is known about whether larger testes have more sperm-producing tissue or produce sperm at a faster rate. Using a comparative approach in New World Blackbirds (Icteridae), we found (1) a strong link between testis histology and sperm length, suggesting selection on testis architecture through selection on sperm size, and (2) that species under intense sperm competition had a greater proportion of sperm-producing tissue within their testes. These results support the prediction that sperm competition fosters adaptations in reproductive organs that extend beyond testes size, and raise questions about the trade-offs influencing reproductive investment.