Moving in two worlds: aquatic and terrestrial locomotion in sea snakes (Laticauda colubrina,Laticaudidae)

Yellow‐lipped sea kraits (Laticauda colubrina) are amphibious in their habits. We measured their locomotor speeds in water and on land to investigate two topics: (1) to what degree have adaptations to increase swimming speed (paddle‐like tail etc.) reduced terrestrial locomotor ability in sea kraits?; and (2) do a sea krait’s sex and body size influence its locomotor ability in these two habitats, as might be expected from the fact that different age and sex classes of sea kraits use the marine and terrestrial environments in different ways? To estimate ancestral states for locomotor performance, we measured speeds of three species of Australian terrestrial elapids that spend part of their time foraging in water. The evolutionary modifications of Laticauda for marine life have enhanced their swimming speeds by about 60%, but decreased their terrestrial locomotor speed by about 80%. Larger snakes moved faster than smaller individuals in absolute terms but were slower in terms of body lengths travelled per second, especially on land. Male sea kraits were faster than females (independent of the body‐size effect), especially on land. Prey items in the gut reduced locomotor speeds both on land and in water. Proteroglyphous snakes may offer exceptional opportunities to study phylogenetic shifts in locomotor ability, because (1) they display multiple independent evolutionary shifts from terrestrial to aquatic habits, and (2) one proteroglyph lineage (the laticaudids) displays considerable intraspecific and interspecific diversity in terms of the degree to which they use terrestrial vs. aquatic habitats.     

Interspecific body size differentiation in species assemblages of the carabid subgenus Ohomopterus in Japan

Geographic variation and interspecific differentiation in body size (body length) were analyzed for 15 species of the carabid subgenus Ohomopterus (genus Carabus; Coleoptera, Carabidae) in Japan. Local species assemblages of this subgenus consist of up to 5 species of different size classes. These beetles exhibited sexual dimorphism in body size where females are larger than males, except Carabus uenoi, in which the male and female sizes were equivalent, possibly because of the exaggerated male genitalia. In 9 of 15 species, there was a positive correlation between mean body size and annual mean temperature of habitat, representing the converse of Bergmann's rule. However, in some cases this correlation does not hold over the range of a species because of regional differences. When allopatric and sympatric populations were compared, allopatric populations of Carabus albrechti and C. japonicus had larger bodies than sympatric populations. These intraspecific differences may have resulted from character displacement. In each local assemblage with 2 or more species, there was little interspecific overlap of body size, although the body size ratio between two species with adjacent body sizes seldom showed strict constancy. The mean size ratio between 2 adjacent species in an assemblage was reduced with the number of species, whereas the size ratio of the largest to smallest species in an assemblage increased with the number of species (i.e., the expansion of body size range). These results indicate that the body size of Ohomopterus species may have evolved in response to both climatic conditions and interspecific interactions. Because each species or species group represents the same size class over the distribution range and similar-sized species are parapatric or allopatric, the interspecific segregation in body size in local assemblages may have resulted mainly from a size assortment process during colonization. Received: June 8, 2000 / Accepted: October 10, 2000     

Microtrichs of amphipod crustacea. Morphology and distribution

Microscopical cuticular structures of four amphipods, presumed to be sensory in function are described, using a scanning electron microscope. Gammarus sp, Liljeborgia sp and Orchomene sp showed very small peg‐like microtrichs emerging from cuticular pits of less than one micron in diameter. Single units, facing distally, were ranged in rows, several rows occupying a single integumental polygon. Density of rows and number of units per row were always higher in central areas. The distribution, organization and relationships with larger sensilla, suggest that microtrichs are specialized in detection of slow or weak water currents. Orchestia showed short sensilla coeloconica. The difference between this species and the other three may be related to its different habitat, which is terrestrial.     

ADAPTATIONS OF TERRESTRIAL ARTHROPODS TO THE ALPINE ENVIRONMENT

1. The climate changes drastically above the timberline. Diverse adaptations have been evolved by insects and other terrestrial arthropods to survive the alpine environment. The fitness of each species depends on a combination of different factors in accordance with their special habitats. 2. Morphological adaptations such as reduced body-size, are known from a number of alpine insects, increasing their possibility to find sheltered microhabitats. Selection for reduced body size in Andean Phulia spp. butterflies is probably a result of their rigorous environment. Wing atrophy, which is also known in insects from other extreme environments, is widespread in alpine species. In several terrestrial arthropods the absorption of solar radiation is increased by melanism. Increased pubescence, protecting against the loss of heat, is known in alpine butterflies and bumblebees. 3. Several behavioural adaptations are described. Thermoregulatory behaviour is important in many species to raise their body temperatures. Alpine butterflies orient the dark basis of their wings perpendicular to the rays of the sun. Body temperatures of 30 °C may be required for flight. To increase their activities many alpine terrestrial arthropods seek warmer microhabitats in the vegetation and under rocks. The adaptive advantage of nocturnal activity as observed in several species, may be to maintain the water balance or to avoid predation. 4. Tropical alpine terrestrial arthropods are faced with special problems. The large diel temperature fluctuations require cold-hardiness during the night and tolerance to heat during the day. Many species seek sheltered microhabitats under rocks and in vegetation. 5. Due to low precipitation and high evaporation rates many mountain areas are extremely dry. High resistance to desiccation may be very important to alpine species, and in particular to tropical species. Rates of water loss at low relative humidities are comparable to those of desert arthropods. 6. As an adaptation to the cold alpine summers several species of terrestrial arthropods require more than one year to complete their life-cycles. Special to these species is their adaptation to low temperatures in two or more overwintering stages. In spite of their cold surroundings several species have univoltine life cycles, frequently combined with highly specialized adaptations. Increased metabolic rates as a compensation to low temperatures may be widespread in alpine species, but few data are available. 7. Cold tolerance is of particular importance in temperature alpine species. Winter survival in Collembola and Acari depends on supercooling. Great seasonal variations have been observed in a number of species. Freezing tolerance is also known from alpine insects, e.g. in some species of beetles. At high latitudes alpine species must endure periods of up to eight or nine month at low temperatures during hibernation. Anaerobiosis is known from species that are enclosed in ice, with lactate as the main end product of metabolism.     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

Is sexual body shape dimorphism consistent in aquatic and terrestrial chelonians?

Comparisons between aquatic and terrestrial species provide an opportunity to examine how sex-specific adaptations interact with the environment to influence body shape. In terrestrial female tortoises, selection for fecundity favors the development of a large internal abdominal cavity to accommodate the clutch; in conspecific males, sexual selection favors mobility with large openings in the shell. To examine to what extent such trends apply in aquatic chelonians we compared the body shape of males and females of two aquatic turtles (Chelodina colliei and Mauremys leprosa). In both species, females were larger than males. When controlled for body size, females exhibited a greater relative internal volume and a higher body condition index than males; both traits potentially correlate positively with fecundity. Males were more streamlined (hydrodynamic), and exhibited larger openings in the shell providing more space to move their longer limbs; such traits probably improve mobility and copulation ability (the males chase and grab the female for copulation). Overall, although the specific constraints imposed by terrestrial and aquatic locomotion shape the morphology of chelonians differently (aquatic turtles were flatter, hence more hydrodynamic than terrestrial tortoises), the direction for sexual shape dimorphism remained unaffected. Our main conclusion is that the direction of sexual shape dimorphism is probably more consistent than sexual size dimorphism in the animal kingdom.     

The evolution of sexual size dimorphism in cottontail rabbits (Sylvilagus,Leporidae)

In mammals, ‘female‐biased’ sexual size dimorphism (SSD), in which females are larger than males, is uncommon. In the present study, we examined Sylvilagus, a purported case of female‐biased SSD, for evolutionary correlations among species between SSD, body‐size, and life‐history variables. We find that: (1) although most species are female‐biased, the degree and direction of SSD vary more than was previously recognized and (2) the degree of SSD decreases with increasing body size. Hence, Sylvilagus provides a new example, unusual for a female‐biased taxon, in which allometry for SSD is consistent with ‘Rensch's Rule’. As a corollary to Rensch's Rule, we observe that changes in SSD in Sylvilagus are typically associated with larger, more significant changes in males than females. Female‐biased SSD could be produced by selection for larger females, smaller males, or both. Although larger female size may be related to high fecundity and the extremely rapid fetal and neonatal growth in Sylvilagus, we find little evidence for a correlation between SSD and various fecundity‐related traits in among‐species comparisons. Smaller male size may confer greater reproductive success through greater mobility and reduced energetic requirements. We propose that a suite of traits (female dispersion, large male home ranges, reduced aggression, and a promiscuous mating system) has favoured smaller males and thus influenced the evolution of SSD in cottontails. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 141–156.     

Granivory in terrestrial isopods

Abstract Granivory (seed feeding) evolved in many animal groups. Field observations hint at the existence of granivory in terrestrial isopods (Crustacea: Isopoda: Oniscidea), for which it was previously unknown. In this paper granivory in terrestrial isopods is addressed for the first time, focusing on (i) seed acceptance in the presence of plant litter and (ii) size as a constraint for acceptance and consumption. In a laboratory choice experiment, Armadillidium vulgare consumed seeds of Capsella bursa‐pastoris and Poa annua when plant litter was present. In a no‐choice experiment, seeds of seven plant species were offered to four isopod species giving 13 combinations in total [A. vulgare (seven species of seeds), Oniscus asellus (two), Porcellio scaber (two), and Porcellionides pruinosus (two)]. The tested isopods differed in their acceptance (proportion of individuals consuming seeds) and consumption (both number and amount of seeds eaten) of seed species. Size as a constraint was demonstrated in A. vulgare offered Cirsium arvense seed, since the probability that this large seed was eaten increased with body size of the isopod. In the other 10 seed–isopod pairs, seed consumption increased linearly with isopod body size. Granivory is thus widespread in terrestrial isopods, although the tendency to eat seeds differs between species.     

Ontogeny and the evolution of adult body size dimorphism in apes

This analysis investigates the ontogeny of body size dimorphism in apes. The processes that lead to adult body size dimorphism are illustrated and described. Potential covariation between ontogenetic processes and socioecological variables is evaluated. Mixed-longitudinal growth data from 395 captive individuals (representing Hylobates lar [gibbon], Hylobates syndactylus [siamang], Pongo pygmaeus [orangutan], Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) form the basis of this study. Results illustrate heterogeneity in the growth processes that produce ape dimorphism. Hylobatids show no sexual differentiation in body weight growth. Adult body size dimorphism in Pongo can be largely attributed to indeterminate male growth. Dimorphism in African apes is produced by two different ontogenetic processes. Both pygmy chimpanzees (Pan paniscus) and gorillas (Gorilla gorilla) become dimorphic primarily through bimaturism (sex differences in duration of growth). In contrast, sex differences in rate of growth account for the majority of dimorphism in common chimpanzees (Pan troglodytes). Diversity in the ontogenetic pathways that produce adult body size dimorphism may be related to multiple evolutionary causes of dimorphism. The lack of sex differences in hylobatid growth is consistent with a monogamous social organization. Adult dimorphism in Pongo can be attributed to sexual selection for indeterminate male growth. Interpretation of dimorphism in African apes is complicated because factors that influence female ontogeny have a substantial effect on the resultant adult dimorphism. Sexual selection for prolonged male growth in gorillas may also increase bimaturism relative to common chimpanzees. Variation in female growth is hypothesized to covary with foraging adaptations and with differences in female competition that result from these foraging adaptations. Variation in male growth probably corresponds to variation in level of sexual selection. © 1995 Wiley-Liss, Inc.     

Different growth responses to temperature and resource limitation in three fly species with similar life histories

Growth responses to temperature and resource limitation in three dipteran species with similar life histories were compared. With respect to current life history theory, two points are raised. First, growth rate in real time increased steeply with temperature in all species, following the standard pattern. However, when expressed in physiological time growth rate increased as temperature decreased in the yellow dung fly Scathophaga stercoraria, remained approximately constant in Sepsis cynipsea, and increased in Drosophila melanogaster. These responses can be understood as adaptations to climate and seasonality. It is concluded that some patterns of adaptation may be more easily interpreted if, and some may even go undetected unless, they are analysed in physiological time. Second, a decrease in body size, development rate and growth rate when resources are limited is believed to be nearly universal and generally predicted by life history models. Despite their similar life histories, the three species investigated showed qualitatively different growth responses to larval food shortage. At unlimited resources, yellow dung flies showed the fastest initial larval body mass gain per unit time, while those of S. cynipsea and D. melanogaster were lower and about equal. The period of no body mass gain at the end of larval development was longest in S. stercoraria and shortest in S. cynipsea. When facing resource limitation, S. stercoraria emerged smaller but earlier (thus nearly maintaining their growth rate), S. cynipsea smaller after the same development period, and D. melanogaster smaller and later (showing reduced and much reduced growth, respectively). It is concluded that whether growth really slows when resources are limited depends on the precise ecological circumstances of the species in question. More refined models, particularly those where mortality costs are independent of time, and more experiments are necessary to account for the variation in growth and size and age at maturity present in nature. This revised version was published online in July 2006 with corrections to the Cover Date.     

Fine structure of the midgut posterior caeca in the crustacean Orchestia in intermolt: Recognition of two distinct segments

This study carried out on the posterior caeca of Orchestia in intermolt by means of light and electron microscopy shows that the diverticula of the midgut consist of two segments which are different from an anatomical point of view. The distal segment is in close relationship to the dorsal blood vessel, whereas the proximal segment, twice as long as the distal one, only touches the haemocoel. The cells of the distal segment are characterized by a brush border, some apical extrusions, a great number of ribosomes, rough endoplasmic reticulum, often associated with the mitochondria, the matrix of which is clear, high activity of the Golgi complexes, and a great development of extracellular channels. All these features indicate an activity in synthesizing proteins and transport. In the proximal segment, the cells are characterized by a striated border, reduced intercellular space, and especially by a great development of the smooth endoplasmic reticulum sometimes associated with mitochondria having a dense matrix. These diverse features indicate absorption ion and water transport. From an ultrastructural point of view, the posterior caeca of Orchestia cannot be considered homologous to the Malpighian tubules. Whereas during molting the posterior caeca of Orchestia are sites of calcium storage, during intermolt they are probably involved in the processes of water and mineral regulation and excretion.     

Cell lineage of the midline cells in the amphipod crustacean Orchestia cavimana (Crustacea, Malacostraca) during formation and separation of the germ band

 Cell lineages of identified midline cells were traced in the amphipod Orchestia cavimana (Crustacea, Malacostraca) by in vivo labelling. Midline cells are a common phenomenon in the germ band of crustaceans and insects. Studies in midline cells of Drosophila showed an origin from separate, paired anlagen and a differentiation into three types of cells. The in vivo labelling of midline cells of Orchestia demonstrates that they originate from the same material as the neural and epidermal ectoderm, divide in a stereotyped cell division pattern and give rise to at least two different types of cells. During the following evolutionarily derived mode of germ band elongation in Orchestia, a morphogenetic process is intercalated that separates germ band halves. On the level of single cells, it can be shown that midline cells are the only ectodermal cells that bridge the large distance between the separated parts. The cells are stretched extensively but do not proliferate. Comparing the midline cells of Orchestia with non-malacostracan crustaceans and insects, the results favour the hypothesis that midline cells are a distinct population of cells homologous in crustaceans and insects. Received: 24 July 1998 / Accepted: 13 October 1998     

Eusynaptomyces benjaminii,spec. nova, (Ascomycetes,Laboulbeniales) und seine Anpassungen an das Fortpflanzungsverhalten seines WirtesEnochrus testaceus (Coleoptera,Hydrophilidae)

Eusynaptomyces benjaminii is described as a new species of the ectoparasiticLaboulbeniales (Ascomycetes). It exists only on two very restricted areas of the body (= position specifity) of its hostEnochrus testaceus (F.) (Coleoptera, Hydrophilidae): on the claws of the right fore-leg and on the lower side of the frontal border of the pronotum. In these two habitatsEu. benjaminii develops two extremely different growth-forms. Male and female hosts are parasitized on somewhat different parts of their body. This can be explained by their mating behaviour. The growth-forms ofEu. benjaminii are so different that one ignorant of the biology of hosts and parasites, might regard them as members of different species or even genera. They are to be interpreted as adaptations of one species to growth positions and mating behaviour of the host. There is no sex-of-host specifity as assumed by certain authors for several species of theLaboulbeniales.

     

Reproductive allocation and reproductive ecology of seven species of Diptera

1. Variation in resource allocation to egg size and number was investigated in seven sympatric species of Piophilidae that oviposit on carcasses or discarded cervid antlers: Liopiophila varipes (Meigen), Prochyliza xanthostoma Walker, Protopiophila latipes (Meigen), Protopiophila litigata Bonduriansky, Stearibia nigriceps (Meigen), and two unidentified species of Parapiophila McAlpine. 2. Following optimal reproductive allocation theory, relatively larger, fewer eggs were expected in (1) species that oviposit on antlers, where larvae probably experience lower risk of predation and greater competition than larvae in carcasses, and (2) species with aggressive males and male-biased sex ratios on the oviposition substrate, where risk of injury during oviposition may have favoured females laying fewer eggs. 3. Variation in reproductive allocation strategies could not be explained by known differences in larval or adult environment, but congeneric species clustered by reproductive allocation patterns. The Parapiophila species produced larger, fewer eggs than the other species, and egg number increased slowly with body size. The Protopiophila species did not deviate from expected egg sizes and numbers, and egg number increased steeply with body size. 4. An interspecific egg size–egg number trade-off resulted in a tight linear scaling of ovary volume to body size, suggesting common physiological constraints on relative ovary mass. 5. Within each species, egg size was nearly constant whereas egg number increased with female body size, suggesting species-specific stabilising selection on egg size.     

Morphological adaptations to arboreal habitats and heart position in species of the neotropical whipsnakes genus Chironius

The evolution of arboreality in snakes is accompanied by modifications that are remarkably similar across species. Gravity is one of the most important selective agents, and arboreal snakes present adaptations to circumvent the gradient of pressure, including modifications on heart position (HP) and body slenderness (BS). However, the degree to which different life‐history traits influence the cardiovascular system of snakes remains unclear. Here, we used an ecological and a phylogenetic approach to explore the relationship between habitat, HP, BS, and heart size (HS) in five species of the neotropical whipsnakes genus Chironius that occupy terrestrial, semiarboreal, and arboreal habits. Our ecological comparison indicated that the arboreal species have the most posterior‐positioned heart, the most slender body, and the smallest HS, whereas the terrestrial representative of the group exhibited the most anterior heart, the less flattened body, and the largest HS. After removing the phylogenetic effect, we found no difference in HP and BS between terrestrial and arboreal species. Habitat only differed when contrasting with HS. Body slenderness and HS were correlated with HP. Our results suggest that different restrictions, such as anatomical constraints, behavior, and phylogenetic inertia, may be important for the studied species.     

The interaction of season length and development time alters size at maturity

An end-of-season penalty, with late-maturing individuals being smaller than early-maturing individuals, has been observed in a variety of univoltine terrestrial arthropods. The current study extends these observations, utilizing multiple populations of a single sexually dimorphic species to examine the ecological correlates and fitness consequences of late maturation at a small size. The orb-weaving spider, Nephila clavipes, inhabits a broad range of habitats that vary from mild to strong seasonality. Because males mature several instars earlier than females, they can reach maturity much earlier in the growing season. Within a cohort, I found that female size at maturity was negatively correlated with timing of maturation in strongly seasonal sites. At a less seasonal site, there was no correlation between female size and timing of maturation within a cohort. In most populations studied, male size was not correlated with the timing of maturation within a cohort. Within populations in strongly seasonal sites, late-maturing females had reduced fecundity. The probability of copulation, survivorship from maturity to first clutch, clutch size relative to female size, and the number of possible clutches were all reduced with delayed maturation. The probability of pre-reproductive death for late-maturing females was strongly affected by stochasticity in the timing of the end of the growing season. Received: 30 December 1998 / Accepted: 1 September 1999     

Host body size as a factor determining the egg complement of Strepsiptera,an insect parasite

The egg complement (total number of eggs produced by a single female) differs greatly among the species of Strepsiptera. The maximum is found in Stichotrema dallatorreanum (750,000 eggs), and the minimum in Triozocera minor (984 eggs). Based on the egg complement of 31 species in 11 genera, the following conclusions were drawn: (1) The egg complement is generally smaller in those species whose hosts gregariously cohabit in a very limited area, or are distinct flower-visitors, compared with those whose hosts display the above two traits weakly; (2) The egg complement is determined by the size of the maternal body. The size of female strepsipterans is reduced when they parasitize smaller host such as males and workers, as compared with those that parasitize larger host such as females and queens; likewise, the size of the strepsipterans becomes larger on increase in size of hosts, showing that their egg complements are principally determined by the size of host species; (3) The increase in the egg complement is compensated for by the reduction in egg size. The relative egg size (length of the firstinstar larva/length of maternal body) is conspicuously reduced according to an increase in the size of the female strepsipterans.     

Consequences of competition on the reproduction and mortality of three species of terrestrial slugs

The interaction of three species of terrestrial slugs was studied in six field cages containing monocultures and 2-way combinations. A. columbianus and A. ater were non-aggressive species and L. maximus was highly aggressive during the summer. All three species were similar in body design and resource utilization. Reproduction (egg size, eggs/batch, number of batches, wet-weight productivity) and mortality were monitored through the 1977 season. L. maximus drastically reduced reproductive success of the non-aggressive slugs and increased their mortality. There were also significant interactions between the non-aggressive species. Compared to other invertebrates, the life history characteristics of A. columbianus are strongly K-selected and probably evolved through intraspecific exploitative competition in predictable, stable forest habitats. A. ater is best classified as a refuging r-selected species adapted to less predictable, unstable field environments. L. maximus appears to be α-selected and its overall design is suited to density-dependent regulation in fields.     

Visual ecology of Indian carpenter bees I: Light intensities and flight activity

Bees are mostly active during the daytime, but nocturnality has been reported in some bee families. We studied temporal flight activity in three species of carpenter bees (genus Xylocopa) in relation to light intensities. X. leucothorax is diurnal, X. tenuiscapa is largely diurnal being only occasionally crepuscular, while X. tranquebarica is truly nocturnal. Occasional forays into dim light by X. tenuiscapa are likely to be due to the availability of richly rewarding Heterophragma quadriloculare (Bignoniaceae) flowers, which open at night. X. tranquebarica can fly even during the moonless parts of nights when light intensities were lower than 10⁻⁵ cd m⁻², which makes this species the only truly nocturnal bee known so far. Other known dim-light species fly during crepuscular or moonlit periods. We compare eye and body sizes with other known diurnal and dim-light bees. We conclude that while extremely large ocellar diameters, large eye size:body size ratio, large number of ommatidia and large ommatidial diameters are all adaptations to dim-light foraging, these alone do not sufficiently explain the flights of X. tranquebarica in extremely dim light. We hypothesise that additional adaptations must confer extreme nocturnality in X. tranquebarica.