Chemical polymorphism in male femoral gland secretions matches polymorphic coloration in common wall lizards (Podarcis muralis)

Previous studies showed that common wall lizards (Podarcis muralis) are polymorphic in colour, both sexes showing three main ventral morphs (white, yellow and red) within the same population and that the three morphs correlate with many life-history traits, including a positive assortative mating according to colour. Chemical communication plays a key role in intra-specific recognition and in social organization of lizards; thus chemical cues might be involved in morph recognition and mate choice. We used gas chromatography–mass spectrometry (GC–MS) to investigate possible differences in the lipophilic fraction of femoral gland secretions between size/age classes and to explore whether chemical secretions match male colour morphs. As expected, most males shared the same compounds, but smaller males showed significantly higher proportions of aldehydes, alcohols and ketones and significantly lower proportions of tocopherols than larger males. Interestingly, inter-morph differences in the proportion of some compounds (especially tocopherols and furanones) matched ventral colour polymorphism. Pairwise comparisons showed that white lizards had significantly different chemical profiles than both the yellow and red ones, whereas differences between yellow and red males were only marginal. A further canonical analysis of principal coordinates correctly classified 67.2 % on average of the chemical profiles according to colour morph (white 85.0 %, red 60.9 %, yellow 57.1 %). We hypothesized that chemical differences associated with colour polymorphism may play a central role in intra-specific communication and even in sexual selection, allowing individuals to choose their partners according to their age, and more interestingly according to their colour morph, in a non-random mating population system.     

Chemosensory discrimination of familiar and unfamiliar conspecifics by lizards: implications of field spatial relationships between males

The ability of territorial lizards to discriminate between scents of neighbors and non-neighbors might contribute to decreasing the costs of aggressive interactions. To test this hypothesis, we conducted a field study to analyze the spatial relationships between male Iberian rock-lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether male lizards can use chemical cues to discriminate between familiar conspecific males (those whose home ranges overlapped) and unfamiliar conspecific males (those whose home ranges did not overlap, and whose home range centers were at least 50 m apart). Differences in tongue-flick rates in the presence of chemical cues suggested that male L. monticola discriminated between odors of familiar and unfamiliar males. The behavioral responses were also dependent on relative differences in body size between the responding male and the unfamiliar male that donated the scent: There was a significant negative correlation between tongue-flick rates emitted in cages of unfamiliar males and the body size differences between males. In contrast, when the donor of the scent was a familiar male, the tongue-flick rate was not dependent on body size differences. These results are compatible with individual discrimination through chemical cues in male L. monticola.     

Environmental and genetic determinants of the male forceps length dimorphism in the European earwig Forficula auricularia L.

Male dimorphisms are particularly conspicuous examples of alternative reproductive strategies. The male forceps length dimorphism in the European earwig Forficula auricularia has long been considered an example of a status- (body size) dependent male dimorphism. In this paper, I test three hypotheses relating to the dimorphism of F. auricularia. First, that the dimorphism is status dependent and determined by nutrition. Second, that the dimorphism is a density-dependent adaptation. Third, that there is a genetic basis to population differences in morph frequency seen in the field. These hypotheses were tested by rearing two populations in a split-family rearing design with two diets and two densities. Populations of male earwigs reared in the common garden differed in forceps length and relative forceps length. The populations also differed in the morph frequencies, with 40 versus 26% long-forceped males. These results confirm the notion that there is a genotype-by-environment interaction that determines the morph frequency in a population. There were only minor effects of density on male forceps length and no influence of density on the male dimorphism. In accordance with the hypothesis that the morphs are status-dependent alternatives, large-forceped males only arose on the high-protein diet that produced earwigs of a large body size. However, not all large males produced the long-forceped phenotype. I put forward an extension of the status-dependent dimorphism model that may account for the pattern of forceps dimorphism in this species. Received: 18 November 1998 / Received in revised form: 14 May 1999 / Accepted: 25 July 1999     

Receiver sex differences in visual response to dynamic motion signals in Sceloporus lizards

Visual signal properties often vary greatly between and within individuals in a variety of social contexts. While it is widely known that visual displays emitted by senders can exhibit great variation in efficacy and content, far less is understood whether and how receivers vary in the ability to respond to variability in signal properties, such as motion. Here, we tested for receiver sex differences in visual response latency to motion signals in Sceloporus undulatus lizards. We used a moving robotic lizard model as a visual stimulus to assay response latency in male and female lizards. We measured visual reaction times to slow and fast, up-and-down motions, characteristic of territorial and courtship male motion displays, respectively. We found sex differences in response latency to the two different displays. Specifically, male lizards were faster than females at responding to slow motion produced by the robotic lizard, while female lizards were faster than males at responding to fast motion. These results demonstrate that dynamic visual signals that vary temporally under different social contexts can differ in eliciting a visual response from each sex. Our study highlights that physical differences in dynamic and complex visual signals exhibited during different social contexts (i.e., territorial and courtship contexts) can closely match sex differences in visual responses.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

Ecological and physiological differences between two colour morphs of the coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus) is a ubiquitous branching coral found throughout the Indo-Pacific region. Like many other species of coral, P. damicornis displays a large range of morphologies. At One Tree Island, it occurs as two distinct morphs that are easily distinguished by the presence or absence of pink pigmentation. The two colour morphs of P. damicornis were found to differ in their distribution and abundance in the One Tree Island Lagoon. The brown morph was more abudant than the pink morph in the shallows (<1 m),whereas the pink morph was more abundant at deeper sites (>3 m). The two morphs also differed physiologically. The brown morph tended to have a greater calcification rate than the pink morph, regardless of environmental conditions. However, the difference in the calcification rate between the two morphs became non-significant under shaded conditions (5% full sunlight), indicating some degree of physiological plasticity of the morphs. The pink colour in P. damicornis was due to a hydrophilic pigment with a major peak absorbance at 560 nm. The expression of pink pigment had both genetic and phenotypic components. The brown morph has a reduced genetic capacity to express the pigment relative to the pink morph. On the other hand, pigment expression could be induced by light in the pink morph. Although genetic differences ultimately determine the differences between the two morphs of P. damicornis, the extent of pigment expression is under some degree of environmental influence.     

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the female's perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.     

Differential breeding investment in bridled and non-bridled common guillemots (Uria aalge): morph of the partner matters

Polymorphism is the coexistence of two or more phenotypically distinct and genetically determinate forms in a population and implies a selective balance between the alternative morphs to permanently exist. Common guillemots Uria aalge occur in two genetically distinct morphs—a non-bridled and a bridled—the latter with white eye rings and a well-defined stripe behind the eyes. In this study, we investigated differences between the morphs with regard to reproductive parameters. We used a detailed family-based sample providing data on mother, father, and chick over three breeding seasons. The mating between morphs was random but pure non-bridled and pure bridled pairs produced smaller chicks at age 15 days than the two mixed pair compositions. Body mass of the adults showed much the same pattern; pure pairs having lower body mass than mixed pairs. There were no differences between the two morphs in reproductive parameters without considering the morph of the partner. This suggests that reproductive decisions in some way not only depend on the morph but also on the tactic of the partner. Different reproductive strategies between morph family groups as found in this study may contribute to the understanding of the existence of a balanced polymorphism in common guillemots. The overall breeding conditions during the years of this study were good. However, over time in a variable environment, we suggest that tactics of different family groups may have different success stabilizing the frequency of non-bridled and bridled birds over time.     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Adaptation,compromise, and constraint: the development,morphometrics, and behavioral basis of a fighter-flier polymorphism in male Hoplothrips karnyi (Insecta: Thysanoptera)

Summary Males of the colonial, wing-polymorphic thrips Hoplothrips karnyi (Hood) fight each other with their forelegs in defense of communal female oviposition areas. In this study, males were reared individually under varying conditions of food deprivation to investigate the developmental cues used in morph determination and the relationships between wing morph, developmental time in each instar, propupal weight, and five adult morphological characters associated with fighting ability and dispersal ability. Males deprived of food for five days midway through the second (final) larval instar had smaller propupal weights and were more likely to develop wings than males deprived of food in the first instar or control males. However, the mean propupal weight of all males that developed wings was not significantly less than that of wingless males. Wing morph of female parents had no measurable effect on this character in the offspring. Wingless males possess relatively larger fore-femora and prothoraces than do winged males, but winged males possess relatively larger pterothoraces (Fig. 1). Behavioral observations of wingless and winged males of similar weight as propupae showed that wingless males won fights and became dominant in oviposition areas. Thus, a trade-off exists between characters associated with male fighting and dispersal ability. The cost of wings, in terms of fore-femora size and prothorax size, increased with propupal weight. Wingless males that developed in the experimental treatment that produced a high proportion of winged males were relatively small in size, and were intermediate in body shape with respect to winged males and other wingless males (Fig. 2). This shape intermediacy indicates that there may be developmental constraints on alternative tactics of resource allocation. Total developmental time varied between wing morphs, but was not correlated with propupal weight or adult morphological characters of winged or wingless males. For wingless males that developed in the treatment that produced a high proportion of winged males, adult morphological characters were negatively correlated with the duration of the second instar. This correlation suggests that the development of small wingless males involves a compromise between the benefits of large adult size and the costs of prolonging the second instar to increase the probability of becoming larger.     

The scent of sun worship: basking experience alters scent mark composition in male lizards

Signals used in female choice should honestly advertise the benefits that males can provide, with direct benefits often argued as being more important than indirect benefits. However, the nature of direct benefits in species without paternal care or nuptial gifts is poorly understood. Previous studies on lizards suggest that females decide where to settle and assumedly who to mate with based on information contained in scent marks from territorial males. Access to high-quality thermal resources is crucial for female reproductive success. Females may therefore be able to detect and exploit thermal-induced variation in the chemical composition of male scent marks when assessing the quality of his territory. We show that the amount of time male wall lizards (Podarcis muralis) are allowed to bask significantly alters the chemical composition of their femoral secretions used in scent marking. The direction of the change is consistent with adaptive plasticity to maintain signalling efficacy under warm conditions that increase evaporation of femoral secretions. The compounds affected by basking experience included those previously associated with male quality or shown to mediate male–male competition in lizards. However, whilst female lizards could discriminate between scent marks of males that had experienced different basking conditions, they did not preferentially associate with the scent from males from high-quality thermal conditions. These results highlight the potential importance of a previously neglected environmental effect on chemical signalling. We suggest thermal effects may have significant consequences for scent-mark composition in variable environments, with potential repercussions on olfactory communication in lizards.     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Experimental evidence of specialised phenotypic roles in a mobbing raptor

Group living is associated with costs but also with potential benefits, such as a decrease in predation risk through, for example, higher defence efficiency. Mobbing is among the most specialised forms of anti-predator strategies involving group defence and has mainly been investigated in passerine birds and some mammals. Variation in the mobbing response has been found in several species according to phenotypic variation such as sex or age. This suggests that there are differential benefits between mobbers, which may have promoted individual specialisation in mobbing behaviour. We studied mobbing behaviour in a communal roosting raptor, the Marsh harrier (Circus aeruginosus), which shows active group defence. Our study population exhibits extreme colour polymorphism, with two colour morphs in males, as well as sexual dichromatism and colour variation with age. We used different decoys, placed at different distances from the roost, to manipulate experimentally the perceived predation risk and to elicit mobbing behaviour. Using the experimental design that maximised mobbing response in harriers, we then focused on the sequence and the specific behaviours involved in recruitment of mobbers, and whether individual investment in terms of defence was associated with phenotypic characteristics of individuals (i.e. sex, age and colour morph). We found that the main behaviour involved in successfully attracting mobbers was alarm calling. We also detected differential individual investment in relation to sex and age, but more importantly, we provide the first evidence for specialised male phenotypic roles during mobbing events, signalled by colour polymorphism: grey males tended not to be involved in mobbing and almost never behaved as recruiters or mobbers, while brown males behaved mainly as recruiter birds. These findings suggest that colour morph may signal the individual’s anti-predatory abilities through different behavioural strategies between males.     

The role of size and aggression in intrasexual male competition in a social lizard species, Egernia whitii

Competition between males is a key component of the agonistic intrasexual interactions that influence resource acquisition, social system dynamics, and ultimately reproductive success. Sexual selection theory predicts that traits that enhance success in intrasexual competition (particularly male–male competition) should be favored. In vertebrates, this often includes body size and aggression, with larger and/or more aggressive males outcompeting smaller or less aggressive conspecifics. The majority of studies consider aggression as a flexible trait which responds to local social or environmental conditions. However, aggression frequently shows considerable within-individual consistency (i.e., individuals have identifiable aggressive behavioral types). Little is known about how such consistency in aggression may influence competition outcomes. We integrated a detailed field study with a laboratory experiment to examine how a male’s aggressive phenotype and his size influence competitive interactions in Egernia whitii, a social lizard species which exhibits strong competition over resources (limited permanent shelter sites and basking sites). Individual aggression and size did not predict competition outcome in the laboratory nor did they predict home range size, overlap, or reproductive success in the field. However, winners of laboratory trial contests maintained consistent aggressive phenotypes while consistency in aggression was lost in losers. We suggest that aggression may be important in other functional contexts, such as parental care, and that alternative traits, such as fighting experience, may be important in determining competition outcome in this species.     

Female lizards discriminate between potential reproductive partners using multiple male traits when territory cues are absent

Female choice can powerfully influence the evolution of male phenotypes. In territorial species, it is challenging to determine the targets of female choice because male traits (e.g., behavior and morphology) are often correlated with territory. We sought to elucidate if and how females specifically evaluate male traits in a territorial species. In this study, we presented female fence lizards, Sceloporus undulatus, with two potential mates to examine mate choice in the absence of territory cues. Females associated more with males possessing better body condition, longer heads, and wider throat badges, and that performed more shudder behavior, which females responded to by approaching shuddering males and performing push-ups. A post hoc decision tree analysis suggests that the strongest predictor of female association was an overall quality index that incorporates all of these traits, rather than individual traits. Male snout–vent length, head width, abdominal badge width, and push-up behavior did not affect female association. Further research on why these traits, which are known to correlate with fitness, do not appear to be used by females when selecting mates would improve our understanding of the evolution of male traits. Our study reveals that females of this territorial species possess the ability to use multiple male traits interactively to make fitness-relevant mate choice decisions in the absence of direct territory cues.     

Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae)

Adult dung beetles (Onthophagus acuminatus) exhibit continuous variation in body size resulting from differential nutritional conditions experienced during larval development. Males of this species have a pair of horns that protrude from the base of the head, and the lengths of these horns are bimodally distributed in natural populations. Males growing larger than a threshold body size develop long horns, and males that do not achieve this size grow only rudimentary horns or no horns at all. Previous studies of other horned beetle species have shown that horned and hornless males often have different types of reproductive behavior. Here I describe the mating behaviors of the two male morphs of O. acuminatus during encounters with females. Females excavate tunnels beneath dung, where they feed, mate and provision eggs. Large, horned males were found to guard entrances to tunnels containing females. These males fought with all other males that attempted to enter these tunnels. In contrast, small, hornless males encountered females by sneaking into tunnels guarded by other males. In many instances, this was accomplished by digging new tunnels that intercepted the guarded tunnels below ground. Side-tunneling behavior allowed sneaking males to enter tunnels beneath the guarding male, and mate with females undetected. Both overall body size and relative horn length significantly affected the outcome of fights over tunnel ownership. These results suggest that alternative reproductive tactics may favor divergence in male horn morphology, with long horns favored in males large enough to guard tunnels, and hornlessness favored in smaller males that adopt the “sneaking” behavioral alternative. Received: 12 October 1996 / Accepted after revision: 8 August 1997     

Male reproductive investment relative to age and flight behaviour in the monandrous butterfly Pararge aegeria

Male reproductive investment may signify a considerable cost to male insects that produce sperm packages or spermatophores. Male butterflies allocate much of their active time budget to mate location, and they may adopt different behavioural strategies to do so. In the speckled wood butterfly (Pararge aegeria L.), males adopt either a territorial wait-and-fight strategy (territorial perching) or a fly-and-search strategy in wider areas (patrolling). In this study, we analysed the impact of male age, male size and male behaviour (i.e. behavioural strategies and levels of activity) on spermatophore investment (i.e. spermatophore mass, number of eupyrene sperm bundles). As predicted, reproductive investment increased with male age and size. Nevertheless, the increase of spermatophore mass with age and the number of eupyrene sperm bundles (i.e. fertile sperm) was stronger in low-activity males compared to active flying males. This suggests that flight activity has a negative impact on male reproductive investment. However, males that were forced to fly in the laboratory produced more eupyrene sperm bundles than resting males. We discuss the potential effects of male–male competition and predation risk on current versus future male reproduction. Males adopting different mate-locating strategies (perching and patrolling) in outdoor cages did not differ in spermatophore traits as was predicted from their very different flight performances. Copulations of territorial perching males took somewhat longer than copulations with non-perching males. There was a significant family effect of spermatophore size and of the expression of male mate-locating strategies suggesting heritable variation. Female traits (i.e. age and size) did not strongly affect spermatophore production. We discuss the results relative to both ultimate and proximate explanations of the complex relationships between butterfly activity, behavioural strategies, age and spermatophore production.     

Lack of innate preference for morph and species identity in mate-searching <Emphasis Type="Italic">Enallagma</Emphasis> damselflies

Insect mate recognition is often viewed as stereotypic, innate, and species-specific. However, male damselflies can learn to identify female-specific color morphs as potential mates. A suite of male mimicry hypotheses assume that heteromorphic females, which differ from males in color pattern, are more easily recognized as “female” and thus lack the inherent, anti-harassment advantage that the more male-like signal provides for andromorphs. Using two measures of male preference, we investigated whether naïve males have a preexisting sensory bias for a given morph color in Enallagma civile, a species that appeared to exhibit extreme plasticity in morph expression across generations within a breeding season. E. civile males raised in the absence of females exhibited no preference for either morph, whereas males raised with one female type exhibited a learned sensory bias for that morph. Male Enallagma also lacked a bias toward conspecific females over a congeneric sister species. In a naturally naïve population of Enallagma ebrium, males reacted sexually to both morphs of Enallagma hageni as often as they did to conspecific females, whose thoracic spectra were nearly identical with those of E. hageni. Moreover, despite the similar thoracic spectra of males and andromorphs, both of which reflected UV, males rarely reacted sexually to other males. Our results falsified implicit assumptions of male mimicry hypotheses, supported learned mate recognition, and suggested a scenario for speciation via sexual conflict.     

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

The hydrozoan coral<Emphasis Type="Italic"> Millepora dichotoma</Emphasis>: speciation or phenotypic plasticity?

This article describes ecological and biological differences between two morphs of the Red Sea fire coral Millepora dichotoma. The species is divided into two main morphs: branching and encrusting, which were found to differ both in color and morphology. Each morph has two or more sub-morphs. A total of 372 M. dichotoma colonies were examined in a census at two study sites in the Gulf of Elat. Colony size and abundance of the two morphs were found to differ significantly between sites. Experimental examination of each morph's morphological plasticity revealed different growth rates and difference in growth plasticity between the branching and the encrusting morph. Most of the fragments from the branching colonies (94%) attached to experimentally placed Plexiglas substrate, compared with much less attachment by the encrusting fragments (11%). The growth form of the branching morph on the Plexiglas switched to encrusting, spreading over and covering the substrate. When the new encrusting colony reached the edge of this substrate, it started to produce tips, and returned to growth in the classic branching form. The encrusting morph did not change its growth form. Following attachment of the original fragments of the branching morph to the substrate, 8.1% of them produced new tips. When the original branches were removed, after converting to encrusting growth form, 19% of the fragments produced new tips. The capsule size of nematocysts of the two morphs was also significantly different (t-test, P<0.05). Molecular data (ITS region) clearly demonstrate that these two M. dichotoma morphs differ considerably. Molecular evidence (srRNA) from the symbiotic zooxanthellae also shows a different pattern of clades in the hosts. The ecological, biological and molecular data thus attest to the two morphs being distinguishable. Contrary to previous reports, we consequently suggest that the two morphs of M. dichotoma found in the Gulf of Elat are actually two distinct species.Communicated by O. Kinne, Oldendorf/Luhe