Copulation Duration Research Articles

Efecto de la duración de cópula en la respuesta ovulatoria y tasa de preñez en alpacas

An experiment was carried out at the Experimental Center La Raya-Universidad Nacional del Altiplano Puno, to determine the effect of copulation duration on ovulation and pregnancy in alpacas. 47 adult females were used (postpartum time ≥ 20 days and presence of a follicle ≥ 7 mm) distributed in three groups, according to copulation time: 20, 35 and 50 min, for G1 (n = 15), G2 (n = 16) and G3 (n = 16), respectively. For the mating, 6 reproducers of proven fertility were used, interrupting the copulation at the established time. Ovulation and pregnancy rates were evaluated on days 7 and 30 post service respectively (by ultrasound and receptivity of the female), the data were analyzed using X2 and simple correspondence. Both the ovulation rate (G1: 73.33%, G2: 81.25% and G3: 81.25%); and pregnancy (G1: 53.33%, G2: 81.25% and G3: 75%) were not different (P> 0.05). Pregnancy rates between G1 and G2 tended to be different (P <0.10), consistent with the simple correspondence analysis. Despite there being no differences in ovulation and pregnancy rates, there was a higher percentage trend in G2 and G3, suggesting that the longer duration of mating would have some positive effect on them, according to the simple correspondence analysis, and the trend of low significance (P <0.10) in favor of pregnancy in G2 compared to G1. In conclusion, there were no significant differences in the effect of copulation duration on ovulation; however, there was a 90% trend for the difference between mating duration of 35 minutes versus 20 minutes (in pregnancy).

Testing potential mechanisms of conspecific sperm precedence in Drosophila pseudoobscura.

Drosophila pseudoobscura females that co-occur with sister species D. persimilis show elevated fertilization by conspecific sperm when they mate with both a heterospecific and a conspecific male. This phenomenon, known as conspecific sperm precedence (CSP), has evolved as a mechanism to avoid maladaptive hybridization with D. persimilis. In this study, we assessed pericopulatory (during mating) and postcopulatory (after mating) traits in crosses with sympatric or allopatric D. pseudoobscura females and conspecific or heterospecific males to evaluate potential mechanisms of CSP in this system. We observed shorter copulation duration in crosses with sympatric females, but found no difference in quantity of sperm transferred or female reproductive tract toxicity between sympatry and allopatry. Our data show some support for the hypothesis that parasperm, a short, sterile sperm morph, can protect fertile eusperm from the D. pseudoobscura female reproductive tract, though it is unclear how this might affect patterns of sperm use in sympatry vs. allopatry. Overall, these results suggest that copulation duration could potentially contribute to the elevated CSP observed in sympatry.

Male age and sexual experience affect male mating behavior in the hawthorn spider mite, Amphitetranychus viennensis.

As is reported, in species with first-male sperm precedence, male age and previous sexual experience play crucial roles in male mating behavior. In the hawthorn spider mite, Amphitetranychus viennensis Zacher, previous studies showed that only females that copulated for the first time could achieve fertilization. Based on this, the effects of male age and mating history on male mate choice and male mate competition were investigated. It was confirmed that males could distinguish virgins from fertilized females but they were unable to discriminate between virgins and unfertilized females. Interestingly, the copulation duration of males mated with fertilized females was much shorter than that of males mated with virgins or unfertilized females. Additionally, for male mating choice, males of all ages and more experienced males preferred 5-day-old virgin females, whereas only less experienced males preferred 1-day-old virgin females. In male-male competition, 3-day-old males were more competitive and obtained more copulations compared with others. Copula duration was closely related to male age. Though no significant differences were observed in mating competition between virgin and mated males, copula duration of males in first copulation was the longest and gradually shortened in subsequent copulations. In all, this investigation demonstrated that male age and sexual experience affected male mate choice and male-male competition, leading to further insight into the influences of male age and sexual experience on the reproductive fitness of both sexes.

Reproductive behaviour in free-ranging crested porcupine Hystrix cristata L., 1758

Previous data on crested porcupine reproduction biology were mostly collected in captivity or semi-captivity due to its elusive, burrowing and mainly nocturnal habits. Between 2017 and 2019 the reproductive behaviour (i.e., intrapair mounting and copulation, birth and parental care) of free-ranging crested porcupine was documented and described. Nightly rhythms of single and multiple mounting occurred throughout the year while only two copulation events were recorded. Three months after both copulation events, the birth of porcupettes (porcupine < 5 kg) was recorded. A stochastic phase characterized by an articulate courtship with distinct behavioural patterns always preceded each mounting event. After the stochastic phase, the presenting of the female evoked by Nose-Quill contact behaviour, occurred in 83.8% (n = 182) of mounting events, while in 16.1% (n = 35) a spontaneous presenting of the female occurred. The average duration of copulation lasted 24 s (SD = 7 s) with 17 thrusting (SD = 5.5 thrusting). Births of porcupettes occurred throughout the year from 1 to 3 times per pair per year and the litter size observed ranged from 1 to 2 porcupettes. The first emerging of porcupettes from burrow occurred at 10–15 days after birth. Parents and sub-adults of the family actively perform parental care and the permanence of sub-adults within the family (i.e., from birth to dispersal) resulted to be at least 1 year. This investigation provides new useful insights on porcupine reproductive behaviour in the wild. Such new knowledge may be useful to the management of porcupines in wild, semiwild and captive condition as well as to delineate the key habitat desiderata of this rodent.

Social and physical environment independently affect oviposition decisions in Drosophila.

In response to environmental stimuli, including variation in the presence of conspecifics, genotypes show highly plastic responses in behavioral and physiological traits influencing reproduction. Although extensively documented in males, such female responses are rather less studied. We expect females to be highly responsive to environmental variation and to differentially allocate resources to increase offspring fitness, given the major contribution of mothers to offspring number, size, and developmental conditions. Using Drosophila melanogaster, we (a) manipulate exposure to conspecific females, which mothers could use to anticipate the number of potential mates and larval density, and; (b) test how this interacts with the spatial distribution of potential oviposition sites, with females from higher densities expected to prefer clustered resources that can support a larger number of larvae. We found that high density females were slower to start copulating and reduced their copulation duration, the opposite effect to that observed in males. There was a parallel, perhaps related, effect on egg production: females previously housed in groups laid fewer eggs than those housed in solitude. Resource patchiness also influenced oviposition behavior: females preferred aggregated substrate, which attracted more females to lay eggs. However, we found no interaction between prior housing conditions and resource patchiness, indicating that females did not perceive the value of different resource distributions differently when exposed to environments that could signal expected levels of larval competition. We show that, although exposure to consexual competition changes copulatory behaviors of females, the distribution of oviposition resources has a greater effect on oviposition decisions.

Utilization of Courtship Behavioral Elements to Understand the Toxicity of Methyl methanesulfonate in &lt;i&gt;Drosophila melanogaster&lt;/i&gt;

A standard, monofunctional alkylating agent, Methyl methanesulfonate, was tested for its effects on courtship behavioral elements in Drosophila melanogaster. Different concentrations ranging from 0.1 to 3 mM were analyzed using the larval feeding technique. When the different combinations of crosses were performed to analyze the effects, the results have revealed that male courtship elements such as orientation, tapping, wing vibration and licking were significantly increased compared to controls. Similar data were observed in case of female behavior. Nonetheless, the courtship behavior interms of fitness such as courtship latency and copulation latency were significantly high and copulation duration was significantly reduced, thus emphasizing the effect on fitness and inturn viability. On par with earlier data on toxicity, the present results also demonstrate the drastic effects of Methyl Methanesulfonate (MMS) on the courtship behavior of D. melanogaster. Thus, it is opined that courtship elements as a parameter can be utilized to understand toxicity and also it can be analyzed within a short period of time.

Mating preference is mating system and relatedness dependent in the ladybird beetle, Menochilus sexmaculatus

AbstractMate choice results from the sensory or behavioural properties that cause individuals to mate with some phenotypes and not others. It can result in direct or indirect (genetic) benefits. Mate choice can also be influenced by various abiotic and biotic factors including the mating system. Here, we investigated how the mating system of the ladybird, Menochilus sexmaculatus, influences mate choice and also affects the time of commencement of mating, copulation duration, fecundity and percent egg viability. We also tested whether mate choice is dynamically affected by relatedness. We found that monandrous and polyandrous females both preferred polyandrous rather than monandrous males. In contrast, males preferred monandrous females over polyandrous females. However, females preferred unrelated monogynous males which were earlier rejected over related polygynous and males preferred unrelated polyandrous females over related monandrous females. Mating system and relatedness also impacted time to commence mating and copulation duration, fecundity and percent egg viability. This will help us to understand precisely why these shifts occur, which will be the subject of future study.

Effects of body size divergence on male mating tactics in the ground beetle Carabus japonicus.

Animal body size is involved in reproduction in various ways. Carabus japonicus exhibits considerable variation in adult body size across geographical locations depending on the larval environment. To investigate the effects of body size divergence on male mating traits, spermatophore deposition and weight, copulation duration, and post-copulatory mounting were observed using male-female pairs from C. japonicus populations with different body sizes. Then, variables with high predictive power on the mating traits were identified from individual characteristics. When the male was slightly smaller than his mate, spermatophore deposition likely succeeded, suggesting that mechanical size-assortative insemination determined male body size. Although male reproductive organ size was positively correlated with male body size, spermatophore weight was not significantly affected by male body size, whereas copulation duration decreased with increasing male body size. Enlarged males, with a high capacity for spermatophore production, could increase paternity by decreasing copulation duration and increasing mating frequency. Such shifts in mating tactics would alter selection pressures of intra- and intersexual interactions (e.g., sperm competition and sexual conflict). Genital dimensions also affected mating traits other than copulatory duration. Thus, ecological heterogeneity has the potential to lead to divergences in sexual traits, such as genital morphology, through body size divergence.

Genital movements are not restricted to spermatozoa transfer in a haplogyne spider

AbstractInternally fertilized animals are characterized by the transfer of their spermatozoa during copulation. However, the duration of copulation is highly variable, which suggests that they may serve for other functions apart from spermatozoa transfer. For example, during copulation, males can stimulate the female by using genitalic movements and/or by positioning their spermatozoa adequately within the female genitalia. In the spider Holocnemus pluchei, males perform strong squeezes and torsion movements with their pedipalps (part of their genitalia actively related to copulation) inside the female genitalia and transfer their spermatozoa during Phase I of copulation. However, the exact moment in which such transfer occurs and its relation to the pedipalp movements are still unknown. Herein, we first have identified the precise moment when the spermatozoa transfer occurs during phase I and its relation to genital movements by interrupting mating couples at the beginning, at middle and at end of this phase. Subsequently, the spermatozoa number remaining in male genitalia and stored in female genitalia were counted. We detected no relation between the number of genital movements and the spermatozoa number present in both, female and male genitalia and body size. As stated above, male genital movements serve another role during copulation in addition to spermatozoa transfer in a context of post‐copulatory sexual selection; most likely, they stimulate the female in a cryptic female choice framework.

Drosophila Corazonin Neurons as a Hub for Regulating Growth, Stress Responses, Ethanol-Related Behaviors, Copulation Persistence and Sexually Dimorphic Reward Pathways.

The neuronal mechanisms by which complex behaviors are coordinated and timed often involve neuropeptidergic regulation of stress and reward pathways. Recent studies of the neuropeptide Corazonin (Crz), a homolog of the mammalian Gonadotrophin Releasing Hormone (GnRH), have suggested its crucial role in the regulation of growth, internal states and behavioral decision making. We focus this review on Crz neurons with the goal to (1) highlight the diverse roles of Crz neuron function, including mechanisms that may be independent of the Crz peptide, (2) emphasize current gaps in knowledge about Crz neuron functions, and (3) propose exciting ideas of novel research directions involving the use of Crz neurons. We describe the different developmental fates of distinct subsets of Crz neurons, including recent findings elucidating the molecular regulation of apoptosis. Crz regulates systemic growth, food intake, stress responses and homeostasis by interacting with the short Neuropeptide F (sNPF) and the steroid hormone ecdysone. Additionally, activation of Crz neurons is shown to be pleasurable by interacting with the Neuropeptide F (NPF) and regulates reward processes such as ejaculation and ethanol-related behaviors in a sexually dimorphic manner. Crz neurons are proposed to be a motivational switch regulating copulation duration using a CaMKII-dependent mechanism described as the first neuronal interval timer lasting longer than a few seconds. Lastly, we propose ideas to use Crz neuron-induced ejaculation to study the effects of fictive mating and sex addiction in flies, as well as to elucidate dimorphic molecular mechanisms underlying reward behaviors and feeding disorders.

Condition-dependent male copulatory courtship and its benefits for females.

Postcopulatory sexual selection has shaped the ornaments used during copulatory courtship. However, we know relatively little about whether these courtship ornaments are costly to produce or whether they provide indirect benefits to females. We used the mealworm beetle, Tenebrio molitor, to explore this. We challenged males using an entomopathogenic fungus and compared their courtship (frequency of leg and antennal contacts to the female), copulation duration, number of eggs laid, and hatching rate against control males. Infected males copulated for longer yet they reduced their leg and antennal contacts compared to control males. However, there was no obvious relation between infection, copulation duration, and courtship with egg production and hatching success. In general, our results indicate that the ornaments used during postcopulatory courtship are condition‐dependent. Moreover, such condition dependence cannot be linked to male fitness.

No water, no mating: Connecting dots from behaviour to pathways.

Insects hold considerable ecological and agricultural importance making it vital to understand the factors impacting their reproductive output. Environmental stressors are examples of such factors which have a substantial and significant influence on insect reproductive fitness. Insects are also ectothermic and small in size which makes them even more susceptible to environmental stresses. The present study assesses the consequence of desiccation on the mating latency and copulations duration in tropical Drosophila melanogaster. We tested flies for these reproductive behavioral parameters at varying body water levels and with whole metabolome analysis in order to gain a further understanding of the physiological response to desiccation. Our results showed that the duration of desiccation is positively correlated with mating latency and mating failure, while having no influence on the copulation duration. The metabolomic analysis revealed three biological pathways highly affected by desiccation: starch and sucrose metabolism, galactose metabolism, and phenylalanine, tyrosine and tryptophan biosynthesis. These results are consistent with carbohydrate metabolism providing an energy source in desiccated flies and also suggests that the phenylalanine biosynthesis pathway plays a role in the reproductive fitness of the flies. Desiccation is a common issue with smaller insects, like Drosophila and other tropical insects, and our findings indicate that this lack of ambient water can immediately and drastically affect the insect reproductive behaviour, which becomes more crucial because of unpredictable and dynamic weather conditions.

Traumatic mating increases anchorage of mating male and reduces female remating duration and fecundity in a scorpionfly species.

Traumatic mating is the male wounding his mate during mating using specialized anatomy. However, why males have evolved to injure their mates during mating remains poorly understood. We studied traumatic mating in Dicerapanorpa magna to determine its effects on male and female fitness. The sharp teeth on male gonostyli penetrate the female genitalia and cause copulatory wounds, and the number of scars on the female genitals is positively related to the number of times females mated. When the injurious teeth were encased with low-temperature wax, preventing their penetration of the female's genitalia during mating, male mating success and copulation duration were reduced significantly, indicating the importance of the teeth in allowing the male to secure copulation, remain in copula and effectively inseminate his mate. The remating experiments showed that traumatic mating had little effect on the female mating refractory period, but significantly reduced female remating duration with subsequent males, probably benefiting the first-mating male with longer copulation duration and transferring more sperm into the female's spermatheca. The copulatory wounds reduced female fecundity, but did not accelerate the timing of egg deposition. This is probably the first report that traumatic mating reduces female remating duration through successive remating experiments in animals. Overall, our results provide evidence that traumatic mating in the scorpionfly helps increase the male's anchoring control during mating and provides him advantage in sperm competition, but at the expense of lowering female fecundity.

Copulatory Signaling and Polygamy of Glassy-Winged Sharpshooters (Hemiptera: Cicadellidae)

AbstractFinding a partner to mate with may be only part of ensuring successful siring of offspring. Females often exhibit cryptic female choice (CFC) during or after copulation, which can influence whose sperm from her multiple partners is chosen for egg fertilization. Known behavioral mechanisms for CFC include assessment of males by their nuptial gifts, duration of copulation, and seminal fluid contents. In this study, the glassy-winged sharpshooter, Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), behaviors during the course of copulation were investigated. Glassy-winged sharpshooter (GWSS) use vibrational communication before copulation occurs. However, little is known about behaviors that occur during and after copulation. Results from this study determined that vibrational signaling also occurs during copulation. Vibrational signals similar to those emitted during precopulatory communication were identified during copulation alongside a new, ‘hum-like’ signal that typically occurred within 10 s after the pair joined in copulation. In addition, results determined the duration of copulation was on average of 15 h, though with a 10-h range (8.5–18.5 h) among observed male–female pairs. Finally, both males and females mated more than once. Collectively, results identified key reproductive parameters required for CFC to occur in GWSS. The study expands on the known animals that use CFC and emphasizes the role that copulatory vibrational communication may play setting the foundations for future more in-depth studies. Understanding of insect behaviors necessary for successful production of offspring is important from an ecological perspective and for development of pest control methods.

Correlated sexual selection on male genitalia, copulatory performance and nuptial gifts in a bushcricket (Orthoptera: Tettigoniidae) indicated by allometric scaling

Abstract We adopt an allometric framework of scaling relationships for comparison between mating-related traits in the middle European bushcricket Roeseliana roeselii (Hagenbach, 1822). Eight characters, covering ontogenetic fitness (size traits; fixed at final moult), male condition (mass traits) and mating motivation (reproductive behaviours), were analysed in unrestricted matings and in matings involving genital manipulation. Shortening the male titillators had no effect on mating-related traits in males. However, titillators, known to be under sexual selection, scale hyperallometrically, with larger males possessing proportionally longer titillators, performing more titillator movements and exhibiting a reduced duration of copulation. Scaling was also hyperallometric for spermatophore mass, with larger males being heavier and transferring heavier nuptial gifts. Both titillator length and spermatophore mass might be condition-dependent indicators, because their variances were nearly twice as large those of body size or body mass. Mass traits were also dynamic, increasing by 11% for male body mass and 17% for spermatophore mass between the first and second matings. Sexual selection by female choice seems to favour larger trait size in the bushcricket R. roeselii, acting in concert on titillator length, intensity of titillator movements and spermatophore mass.

Effects of Delayed Mating on the Reproductive Performance and Longevity of the Fall Armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae).

The fall armyworm, Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae), a severe pest on agricultural crops occurring throughout the tropical Americas, has been reported to occur in China since 2019. To develop novel pest management practices, we studied the effect of delayed mating on the reproductive performance and longevity of S. frugiperda. Delayed mating, progressing from 0 to 7 days, was respectively imposed on both sexes simultaneously, female only, and male only. We demonstrated that delayed mating reduced mating success, number of eggs laid, egg hatch rate, and female oviposition period, while increased copulation duration and longevity. The correlations between the number of delayed days and mating success, number of eggs laid, hatch rate, and oviposition period were all significantly negative irrespective of the sex that was delayed. Meanwhile, there was a positive correlation between delayed days and copulation duration when both sexes were delayed simultaneously or males delayed only. Overall our results indicated that delayed mating in both males and females drastically reduced female reproductive output.

Parents’ living conditions influence offspring fitness and competency

In this study, we used Callosobruchus maculatus as a model insect to assess the idea of whether or not environmental conditions experienced by parents subsequently moderate offspring fitness and competency. We observed that parents’ experiences (sex ratio of parents, restricted access to mates, and honey access) influenced their offspring’s features and did not significantly affect traits such as offspring sex ratio, larval duration, and longevity. However, not only offspring (F1) but also grand-offspring (F2) copulation behaviors, egg number, and hatching percentage were influenced by parents’ experiences. Hence, the offspring of parents living in unfavorable conditions had higher copulation success, higher male mating propensity, and a lower mating latency time in adulthood than the progeny of parents kept under natural or favorable conditions. Interestingly, the offspring of parents with favorable living conditions showed the least tendency to copulate immediately. Even grandparents’ experience could affect descendant behaviors and shorten copulation duration. Adjustments of the parents to their environment were inherited by offspring and grand-offspring to cope better with challenging circumstances and might make the organism more competent. Therefore, the parents’ living conditions have a remarkable impact on how their progeny behave in different situations.

Mating status of coexisting conspecific individuals modulate reproduction in a ladybird, Menochilus sexmaculatus

Abstract The mating status of organisms plays a crucial role in deciphering mating decision and reproductive success of any organisms. Odour or pheromones are known to be perceived by animals through olfaction to locate mates at a distance. In this study, an attempt was made to investigate the effect of the presence of unmated and mated conspecific adults on mating and reproductive parameters of the ladybird beetle, Menochilus sexmaculatus (Fabricius, 1781). To conduct this experiment, an arena was made with the help of two Petri dishes and in such a manner that the experimental adults were only able to perceive the presence of conspecific adults of different mating status in their close vicinity. The results of the study demonstrated an early onset of mating in the presence of males compared to females. Another male in the vicinity poses competition that may force the males to establish genital contact faster. In the absence of any potential rival, copulation duration was highest. Females laid more eggs in the presence of mated females. This may be to increase offspring fitness in the presence of potential competitors for the already mated female. Females laid fewer eggs in the presence of unmated females. Unmated females nearby can be another choice for a male and thus, the male possibly limits the size of the ejaculate containing oviposition stimulants. The current study advocates that mating and reproductive behaviour are modulated according to the perceived surroundings in the form of conspecific adults of different mating status.

Does Larval Rearing Diet Lead to Premating Isolation in Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae)?

Simple SummarySpodoptera litura Fabricius (Lepidoptera: Noctuidae) is a serious polyphagous pest. Most studies focus on the effects of natural hosts on S. litura. However, progressively more laboratory studies S. litura involve feeding the larvae with an artificial diet. We compared the life performance and observed mating choice of S. litura reared on tobacco, Chinese cabbage, and an artificial diet. The results revealed that diet had a significant effect on the duration of each stage of development. In the multiple-choice test with individual males consuming tobacco, Chinese cabbage, or an artificial diet, females fed on the artificial diet preferred to mate with males that were fed on the same diet and rarely mated with males fed on tobacco or Chinese cabbage. We suggest that the diet of S. litura has a potential impact on mate choice and sexual isolation.Host plant preference during the larval stage may help shape not only phenotypic plasticity but also behavioral isolation. We assessed the effects of diet on population parameters and mate choice in Spodoptera litura. We raised larvae fed on tobacco, Chinese cabbage, or an artificial diet, and we observed the shortest developmental time and highest fecundity in individuals fed the artificial diet. However, survival rates were higher for larvae on either of the natural diets. Population parameters including intrinsic rate of increase and finite rate of increase were significantly higher with the artificial diet, but this diet led to a lower mean generation time. Copulation duration, copulation time, and number of eggs reared significantly differed between diets. In terms of mate choice, females on the artificial diet rarely mated with males fed on a natural host. Our results support the hypothesis that different diets may promote behavioral isolation, affecting mating outcomes. Thus, findings for populations fed an artificial diet may not reflect findings for populations in the field.

Life history of the false flower mantid (Harpagomantis tricolor Linnaeus, 1758) (Mantodea: Galinthiadidae) and its distribution in southern Africa

The false flower mantid is the common name for the Mantodea speciesHarpagomantis tricolor(Linnaeus, 1758). This species uses camouflage as a defense mechanism. Limited information (Kaltenbach 1996, 1998) exists on its distribution in southern Africa or about its life history. This species, and Mantodea to an extent, are not usually included in biodiversity studies from this region. The aim of this study was to determine the distribution of this species in southern Africa based on museum collection records and to study the biology ofHarpagomantis tricolorunder captive breeding conditions. The distribution ofHarpagomantisand its morphological variety, i.e.,discolor, were determined utilising the historical insect collection records of seven national museums throughout South Africa. Field collectedH. tricolormales and females were mated and reared under laboratory conditions to record their life history parameters of nymphal duration, oothecae structure, size and incubation duration, adult longevity, and sex ratio. The results of this study indicate that the mean duration of the lifecycle ofH. tricoloris 191.33 ± 37.96 days. All but threeH. tricolorindividuals had five nymphal instars, and the mean duration of the nymphal stage was 140.20 ± 31.03 days. The mean duration of copulation was six hours, while the average incubation period of oothecae was 144.71 ± 9.33 days. These results indicate that oothecae ofH. tricolorprobably overwinter under field conditions and that males of this species have evolved various mechanisms to increase the likelihood of ensuring their own genetic offspring. This study bridges the gap in rudimental research in which Mantodea, in general, has been overlooked and establishes a basis on which ecological interactions, habitat preferences, and imminent threats toH. tricolorcan be established.