Xyleborus Affinis
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''Xyleborus affinis'', the sugarcane shot-hole borer, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
ambrosia beetle Ambrosia beetles are beetles of the weevil subfamilies Scolytinae and Platypodinae (Coleoptera, Curculionidae), which live in nutritional symbiosis with ambrosia fungi. The beetles excavate tunnels in dead or stressed trees into which they introduc ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae. Th ...
. It is found on all continents with woodlands and is one of the most widespread ambrosia beetles internationally, primarily in areas with humid tropical climates like Florida. Like other ambrosia beetles, ''Xyleborus affinis'' is attracted to dead fallen logs in early stages of decay. Due to the presence of ''X. affinis'' on moist timber, this species is mistakenly believed to be the cause of tree death. However, there is insufficient evidence to support their presence as a causal factor. Rather, these beetles are believed to target and accelerate the decay process of trees that are already deteriorated and weak. Similar to other species of ambrosia beetles, ''Xyleborus affinis'' lives in symbiosis with ambrosia fungi and dig tunnels (formally known as galleries) into the
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
of decaying trees where the fungus is then grown. Female eggs are laid in these fungus-lined galleries and use the fungus as a food source. How prominent sexual dimorphism is in this species is highly dependant on temperature conditions during development, with individuals that were reared under high temperatures having less pronounced sexually dimorphic traits than those that were reared under lower temperature conditions.


Subspecies

These two subspecies belong to the species ''Xyleborus affinis'': * ''Xyleborus affinis affinis'' * ''Xyleborus affinis mascarensis'' Eichhoff, 1878


Range

In the United Status, ''Xyleborus affinis'' is found all over the east coast, ranging from Michigan in the north to Florida in the south and even to Texas in the west. This species is also found in regions of South and Central America, such as the Antilles, Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama, and Argentina. The species has also spread into Africa, Asia, Australia, Europe, and the Pacific Islands such as Hawaii. ''Xyleborus affinis'' originates in the American tropics, and is especially abundant in tropical climates like Florida, it currently inhabits the majority of international tropical and subtropical regions. Similar to other beetle species that inoculate wood, ''Xyleborus affinis'' is easily transmitted via wood distribution and international trade.


Characteristics


Adults

This species is yellowish to reddish brown in color and resembles the
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of other ''Xyleborus'' species with its elongated, cylindrical body. ''Xyleborus'' adults are
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, which means that the size, shape, and structure of bodily characteristics differ between males and females. Females are, on average, 2.0–2.7 mm long and are larger than males, which are typically 1.7–2.0 mm long. In contrast to females, males do not have wings, have smaller eyes and antennae, and are less numerous in the population. Although the female to male sex ratio has been reported as 14 to 1 in a study by Roeper et al. (2017), it widely varies and has not yet definitively been identified. This species very closely resembles ''Xyleborus perforans'' Wollaston and ''Xyleborus volvulus'' Fabricius, both of which are found in Florida.''Xyleborus affinis'' is unique based on the downward slant of its abdomen at the end of its
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
. In ''Xyleborus affinis'', the surface of this declivity is dull and nontransparent, whereas it is shiny and smooth in the other two species.


Eggs

The eggs are ivory-colored, ellipsoidal, and shiny and vary in length from 0.6 to 1.0 mm with an average of 0.718 mm. Females lay eggs in groups of two to four along the fungus-lined horizontal galleries that extend horizontally away from the main vertical tunnels. At 29 °C, eggs can be laid anywhere from three to 27 days after introduction into the host log.


Larvae

Larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
are white, have a slightly bent shape, and do not have legs. At 29 °C, larvae take 7–14 days to hatch, whereas they take14–35 days to hatch at 22–24 °C. Larvae are nourished solely from the symbiotic fungus inhabiting the galleries.


Pupae

Pupae A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
are initially white, but they become light brown in color just before they appear as adults. On average, males are 2.0 mm long and shorter than females, which are 2.7 mm long. At 29 °C, it takes larvae 11-23 days to mature into pupae, whereas it takes 21–35 days at 22–24 °C. At 29 °C, it takes adults 18-35 to surface, whereas it takes 27–35 days at 22–24 °C.


Ecology


Hosts

''Xyleborus affinis'' is found in natural woodlands and inhabits damp logs on the ground. Even though this species is one of the most common ambrosia beetles found in wooded regions, its population size is often grossly underestimated. The beetles inoculate large trees in a series of stages, first inhabiting dead
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
and then moving onto the
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
, which is where most eggs are laid. Although ''Xyleborus affinis'' is not particular about choosing the species of its host tree, it is selective based on how deteriorated and damp the fallen log is. These beetles favor large and moist pieces of wood that are early on in the decay process, which is believed to be due to the symbiotic fungus' moisture requirement for survival. As a result, these beetles are especially plentiful in logs partially immersed in water or lying on moist ground.


Symbiosis

The female beetles have a specialized pocket inside their mouths, known as mycangium, where they maintain a population of fungal symbiont. This fungus gets injected inside the galleries the females create, allowing for a mutualistic symbiosis. Until they become adults, larvae will continue to feed on the spore-containing fungi.
Ethanol Ethanol (also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound with the chemical formula . It is an Alcohol (chemistry), alcohol, with its formula also written as , or EtOH, where Et is the ps ...
is the most popularly used chemicals that attracts ambrosia beetles and is used in observational studies. However, ''Xyleborus affinis'' is not strongly drawn toward ethanol and is thus more difficult to monitor.


Reproduction

Like all other ''Xyleborus beetles, Xyleborus affinis'' is
haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
. In this sex-determination system, females develop from fertilized eggs with a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
set of DNA from both their mother and father, whereas males develop from unfertilized eggs and are born
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
, containing DNA from only their mother. Additionally, ''Xyleborus affinis'' is
inbred Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders an ...
. Because males remain confined to their original host tree and their only purpose is to fertilize nearby females, many of these eggs are their sisters. After becoming adults, females will either continue reproducing in the host log or, if the current log is too decomposed, will look for a new host. In contrast to most ambrosia beetles in which each generation seeks a new host, ''Xyleborus affinis'' ability to maintain consecutive populations in the same log is atypical.


Sexual dimorphism


In the genus

The phenomenon of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
within ''Xyleborus'' genus, particularly in terms of size and distinct categorical traits, has been thoroughly established. In research conducted by Ospina-Garcés, Sandra M., et al. (2021), an analysis was carried out to assess the differences in size and shape of the elytra and
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
between sexes. The findings indicated that sex does not affect how size influences shape differences in these body parts, with the degree of shape variation attributable to similar
centroid In mathematics and physics, the centroid, also known as geometric center or center of figure, of a plane figure or solid figure is the arithmetic mean position of all the points in the figure. The same definition extends to any object in n-d ...
size between the sexes and the two structures examined. A significant correlation was identified between the shapes of the pronotum and elytra, suggesting a coordinated development of body shape in these beetles. Furthermore, while both the elytra and pronotum exhibit sexual dimorphism in size and shape, this difference is more pronounced in the shape of the pronotum compared to the elytra. The study also revealed contrasting patterns of shape variation in relation to centroid size for each structure: a negative
allometric Allometry (Ancient Greek "other", "measurement") is the study of the relationship of body size to shape, anatomy, physiology and behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Juli ...
trend was observed for elytra shape from males to females, where as a positive allometric trend was observed for pronotum shape from males to females.


In the species

The findings reveal that structural differences in the elytra and pronotum vary in a coordinated manner. However, the pronotum demonstrates greater variation in shape and therefore a higher degree of sexual dimorphism in both size and shape compared to the elytra. Prior studies, including observations in ''X. affinis,'' have noted sexual size dimorphism in the pronotum, with females displaying a larger size than males when measured using traditional methods of length and width. An additional study by Ospina-Garcés, Sandra M., et al. (2021) detected a notable sexual dimorphism in ''X. affinis,'' with males possessing shorter and wider mandibles compared to females, as well as exhibiting an apophysis on the external contour of the mandible. This dimorphism is thought to be influenced by the species' haplodiploid sex determination system and prevalence of inbreeding, as well as the fact that only females spread out and colonize.


Effect of temperature

The same study evaluated the effect of different temperatures on the development of head size and mandible shape throughout the growth stages of ''X. affinis''. It suggests that the temperature in which the larvae grow contributes to head size variation, where a narrower mandible size range was observed at higher temperatures (width: 0.12–0.40 mm), while lower temperatures allowed for a wider range of mandible sizes (width: 0.11–0.49 mm). It was also found that mandible size variation was linked to changes in larval
instars An instar (, from the Latin ''wikt:instar#Latin, īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each ecdysis, moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the ...
through four distinct ontogenesis stages, regardless of the temperature conditions. Regarding the extent and pattern of sexual dimorphism in mandible shape and size, the study found that the differences between sexes were minimized at higher temperatures, whereas more variable patterns of dimorphism were observed at lower and medium temperatures. The most pronounced disparities in mandible shape between sexes were recorded at 23 °C, and the largest size differences at 26 °C. These findings suggest that extreme temperatures outside optimal temperature ranges (23 °C and 29 °C) may induce phenotypic variations in mandible shape between sexes, potentially affecting their functionality.


Damage

Although the inoculation of ''Xyleborus affinis'' on fallen logs can accelerate the decay process, these beetles typically seek out weak and injured timber that are already deteriorating. As a result, the presence of these beetles alone is typically not believed to cause decomposition. ''Xyleborus affinis'' is gaining attention in the context of plant diseases based on its ability to transmit disease-causing fungi, such as the causal agent of laurel wilt disease, between different plant species. When these beetles burrow into trees in the Laurel family, they transfer the fungal seeds to the trees. These spores proceed to contaminate the xylem bu cutting off the vascular system, leading to wilting and mortality in as quick as weeks to months after inoculation. ''Xyleborus affinis'' can also compromise the structural integrity of freshly cut natural wood more than any other ambrosia species because of its family arrangement and labor division. In most ambrosia beetle species, the mother is solely responsible for burrowing galleries, but in ''Xyleborus affinis'', the daughter females help expand the tunnel system as well. This allows numerous gallery systems to be formed, both superficially and deep within the xylem.


References


Further reading

* * {{Taxonbar, from=Q50389584 Scolytinae Articles created by Qbugbot Beetles described in 1868