0000000000420683
AUTHOR
Katja Tynkkynen
SIMULATING RANGE EXPANSION: MALE SPECIES RECOGNITION AND LOSS OF PREMATING ISOLATION IN DAMSELFLIES
Prolonged periods of allopatry might result in loss of the ability to discriminate against other formerly sympatric species, and can lead to heterospecific matings and hybridization upon secondary contact. Loss of premating isolation during prolonged allopatry can operate in the opposite direction of reinforcement, but has until now been little explored. We investigated how premating isolation between two closely related damselfly species, Calopteryx splendens and C. virgo, might be affected by the expected future northward range expansion of C. splendens into the allopatric zone of C. virgo in northern Scandinavia. We simulated the expected secondary contact by presenting C. splendens fema…
Immunocompetence and resource holding potential in the damselfly, Calopteryx virgo L
It is generally believed that resource holding potential reliably reflects male quality, but empirical evidence showing this is scarce. Here we show that the outcome of male-male competition may predict male immunocompetence in the territorial damselfly, Calopteryx virgo (Odonata: Calopterygidae). We staged contests between 27 pairs of males and found that winners of the contests showed higher immunocompetence, measured as encapsulation response, compared with that of losers. Furthermore, the winners had larger fat reserves. We also collected 29 males that had not been used in staged contests, and found that in these males encapsulation response correlated positively with an individual’s fa…
INTERSPECIFIC AGGRESSION CAUSES NEGATIVE SELECTION ON SEXUAL CHARACTERS
Interspecific aggression originating from mistaken species recognition may cause selection on secondary sexual characters, but this hypothesis has remained untested. Here we report a field experiment designed to test directly whether interspecific aggression causes selection on secondary sexual characters, wing spots, in wild damselfly populations. Males of Calopteryx virgo are more aggressive toward males of C. splendens with large than with small wing spots. This differential interspecific aggression may cause negative selection on wing spot size. Indeed, our results show that directional survival selection on wing spot size of C. splendens males was changed by experimental removal of C. …
Digital Education For All: Multi-University Study of Increasing Competent Student Admissions at Scale
An indubitable way to put learning at scale in practice is to implement Massive Open Online Courses, or MOOCs. When a wide-enough portfolio of them is available, new applications arise. For instance, university admissions in Finland, where this study was conducted, have traditionally been based on students' grades in high school studies, an entrance examination, or a combination of both. A minority of students have been accepted through an open university admission path where students can get a study right if they complete enough university course credits with a high enough grade in a given time frame. In this work, we report results from a multi-university project in which the open univers…
Interspecific territoriality in Calopteryx damselflies: the role of secondary sexual characters
Interspecific territoriality is usually interpreted to result from interspecific interference competition, although it may also originate from mistaken species recognition. In the latter case, it may be based on similarity of secondary sexual characters. In the damselfly Calopteryx splendens, males have pigmented wing spots as a sexual character, and males with the largest spots resemble males of another species, Calopteryx virgo. Probably because of this resemblance, C. virgo males are more aggressive towards large- than small-spotted C. splendens males. We examined whether wing spot size of C. splendens males affects territorial interactions between the species. In a removal experiment, t…
Interspecific interactions and premating reproductive isolation
Interspecific interactions have several evolutionary consequences: for example, two species may compete, hybridize, or behave aggressively towards each other, or there may be predator–prey interactions. One consequence of these interactions is the evolution of premating reproductive isolation between the two species. The most obvious interspecific interaction, which has an effect on reproductive isolation, is the avoidance of hybridization, or, in other words, the reinforcement process. The theory of reinforcement states that when hybridization is maladaptive, selection pressure causes a divergence in female mate preference and/or in male secondary sexual characters. It is often assumed tha…
Alternative reproductive tactics and the propensity of hybridization
One explanation for hybridization between species is the fitness benefits it occasionally confers to the hybridizing individuals. This explanation is possible in species that have evolved alternative male reproductive tactics: individuals with inferior tactics might be more prone to hybridization provided it increases their reproductive success and fitness. Here we experimentally tested whether the propensity of hybridization in the wild depends on male reproductive tactic in Calopteryx splendens damselflies. Counter to our expectation, it was males adopting the superior reproductive tactic (territoriality) that had greatest propensity to hybridize than males adopting the inferior tactics (…
Immunocompetence, developmental stability and wingspot size in the damselflyCalopteryx splendensL.
Calopteryx splendens males exhibit a remarkable variation in wing pigmentation both within and between populations. In this study, we examined whether the wingspots of male C. splendens are related to male quality. We measured the nylon implant encapsulation rate for 85 males and found that males with larger wingspots had a faster encapsulation rate, indicating a better immunocompetence. We also found that the encapsulation rate was positively correlated with the density of haemocytes in the haemolymph. Another measurement of male quality, fluctuating asymmetry of wingspots, correlated negatively with the size of the wingspots. Males with asymmetrical wingspots also had lower encapsulation …
Interspecific interactions and selection on secondary sexual characters in damselflies
Lajienvälinen aggressiivisuus voi selittää immenkorentokoiraiden seksuaaliominaisuuden koossa havaittavaa vaihtelua, toteaa Katja Tynkkynen väitöskirjatyössään. Sekundaarisien seksuaaliominaisuuksien, kuten koiraslintujen kirkkaan värityksen, on tulkittu olevan seurausta seksuaalivalinnasta. Koiraat, joilla on suurempi tai upeampi seksuaaliominaisuus saavat yleensä muita suuremman lisääntymismenestyksen, minkä seurauksena seksuaaliominaisuuksiin kohdistuu positiivista suuntaavaa valintaa. Näin voi tapahtua esimerkiksi naaraan valinnan seurauksena tai koska suuren tai upean seksuaaliominaisuuden omaavat koiraat menestyvät paremmin koiraiden välisessä kilpailussa. Lajiensisäisten vuorovaikutu…
Crystalline wax coverage of the imaginal cuticle inCalopteryx splendens(Odonata: Calopterygidae)
Abstract In this study we use high resolution SEM to describe the diversity of wax crystals and their distribution on different morphological structures in male individuals of Calopteryx splendens. The entire cuticle surface of this damselfly, with the exception of ommatidia and ocelli, is covered with crystalline wax in dimensions from submicron to micron range. It is shown that shape - rod-like, plate like, filamentous, etc. -, size, and density of crystals vary on different surfaces and in individuals of different ages. Additionally, we demonstrate different types of damage to the crystalline wax layer: scratches, compressions, wear, and contamination. The primary function of the wax cry…
The macrophyte, Stratiotes aloides, protects larvae of dragonfly Aeshna viridis against fish predation
Predation could be one force determining which contemporary species occupy a certain habitat. Aeshna viridis is an endangered dragonfly species with a larval distribution strongly associated with lakes where the water plant, water soldier, Stratiotes aloides, occurs. In this study, the larvae were almost exclusively found in patches of S. aloides. To study larval association with S. aloides further, we conducted a series of laboratory experiments. Behavioural experiments indicated that larvae preferred S. aloides. Aeshna viridis larvae were nocturnal and rather inactive. Larvae on S. aloides were less susceptible to predation by the perch, Perca fluviatilis, than larvae on another water pla…
Crystalline wax coverage of the imaginal cuticule in Calopteryx splendens (Odonata: Calopterygidae)
In this study we use high resolution SEM to describe the diversity of wax crystals and their distribution on different morphological structures in male individuals of Calopteryx splendens. The entire cuticle surface of this damselfly, with the exception of ommatidia and ocelli, is covered with crystalline wax in dimensions from submicron to micron range. It is shown that shape – rod-like, plate like, filamentous, etc. –, size, and density of crystals vary on different surfaces and in individuals of different ages. Additionally, we demonstrate different types of damage to the crystalline wax layer: scratches, compressions, wear, and contamination. The primary function of the wax crystalline …
Hybridization in Calopteryx damselflies: the role of males
Females are often considered responsible for hybridization between two species because usually they are the choosier sex and their cooperation is needed for successful copulation. However, males can also be responsible for hybridization, for example in species in which males are able to force copulation. We studied the pattern of hybridization in two congeneric damselfly species, Calopteryx splendens and Calopteryx virgo, and provide evidence that F1 hybrids between the two damselfly species occur in the wild. According to mitochondrial DNA analysis, hybridization is reciprocal: five of seven hybrids were sired by C. splendens and two by C. virgo males. We conducted an experiment that revea…
Interspecific aggression and character displacement in the damselfly Calopteryx splendens
Problems in species recognition are thought to affect the evolution of secondary sexual characters mainly through avoidance of maladaptive hybridization. Another, but much less studied avenue for the evolution of sexual characters due to species recognition problems is through interspecific aggression. In the damselfly, Calopteryx splendens, males have pigmented wing spots as a sexual character. Large-spotted males resemble males of another species, Calopteryx virgo, causing potential problems in species recognition. In this study, we investigate whether there is character displacement in wing spot size and whether interspecific aggression could cause this pattern. We found first that wing …