Research Article |
Corresponding author: Volker Mauss ( volker.mauss@gmx.de ) Academic editor: Jack Neff
© 2016 Volker Mauss, Andreas Müller.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Mauss V, Müller A (2016) Contribution to the bionomics of the pollen wasp Quartinia canariensis Blüthgen, 1958 (Hymenoptera, Vespidae, Masarinae) in Fuerteventura (Canary Islands, Spain). Journal of Hymenoptera Research 50: 1-24. https://doi.org/10.3897/JHR.50.6870
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Quartinia canariensis was recorded from three semidesertic sand habitats in Fuerteventura. All localities were sparsely covered by halophytic vegetation and characterized by large patches of flowering plants of Frankenia laevis (Frankeniaceae). Males and females were exclusively observed to visit flowers of Frankenia laevis. During flower visits the imagines often switched between nectar and pollen uptake. Pollen was consumed directly from the anthers or pollen uptake was indirect with pollen grains gathering on the frons being brushed towards the mouthparts with the fore legs. During nectar uptake the wasps protruded their long proboscis into the nectariferous pockets between the claws of the petals of the Frankenia flowers. Brood cell provisions consisted mainly of pollen from Frankenia but to a small amount also from Polycarpaea (Caryophyllaceae) suggesting that Quartinia canariensis is polylectic with strong preference. Males regularly stood on the ground in the vicinity of Frankenia plants and frequently performed patrol flights along the flowers. Flower visiting females avoided contact with the males and mainly offered resistance against the insertion of the male genitalia. The behavioural sequence during copulation of a species of Quartinia is described for the first time. The nest was a multicellular sub-vertical burrow surmounted by a short turret. The burrow was excavated by the female in friable sandy soil. The walls of the nest were stabilized by silk produced by the nest building female and applied with the mouthparts. Inconsistencies concerning the host-parasite-relationship between Quartinia canariensis and Chrysis atrocomitata established by
Palaearctic, flower associations, mating behaviour, nest construction, silk spinning, Chrysis umbofascialis
The genus Quartinia is a monophyletic subtaxon of the Masarinae (
Quartinia canariensis was described in 1958 by Blüthgen from three females collected by H. Lindberg from Fuerteventura during the Finnish Canary-Expedition. Further material from the Canary Islands, which had been collected by K.M. Guichard between 1964 and 1966, led to the description of the previously unknown male of Q. canariensis by
Published information about the bionomics of Quartinia canariensis is very fragmentary. There are several records of males and females having been observed visiting flowers of Frankenia laevis L. (Frankeniaceae) (
In the present study data concerning flower associations, flower visiting behaviour, nesting and mating of Quartinia canariensis are recorded for the first time, as an example for a Quartinia species from the Palaearctic. A comparison is made with published data on the bionomics of the Afrotropical species of the genus.
Investigations were carried out from 29 March to 5 April 2015 in the vicinity of Costa Calma in Fuerteventura (Canary Islands, Spain). Geographic coordinates (WGS 84) were measured using a Garmin GPS 12. Quartinia canariensis was studied at three localities [I Montaña Pelada, 2 km SW Costa Calma, eastward exposed slope W of a Barranco SW of the road to Los Gorriones, 28°08.325'N 14°14.999'W, 55 m a.s.l.; II SW Costa Calma between “Playa Esmeralda” and “Playa Paraíso” 28°08.677'N 14°14.368'W, 60 m a.s.l.; III Istmo de la Pared, 0.5 km NW Costa Calma 28°10.202'N 14°14.396'W, 90 m a.s.l.]. Most observations were made at locality I.
For all documentations of observations the local time (= Greenwich Mean Time + 1h) was used. Sunrise was approximately at 7h45, sun’s zenith at 14h00 and sunset at 20h15. Time intervals were measured using a digital stop-watch. Observations were made with a close-up binocular (Pentax Papilio 8.5x21) and documented by using a Canon EOS 70D camera with a 180 mm macro lens and a 25 mm extension tube (scale more than 1:1, resolution 20 mega pixel) and macro flash-lights. In addition, in the course of one investigation interval during nest observations, the system was combined with a close-up lens (4 dioptre).
Specimens of all plant species flowering at locality I were collected and preserved dried. The material was placed in the herbarium of the Staatliches Museum für Naturkunde Stuttgart (Herbarium
Activity and behaviour of the female of nest No.1 were observed on 6 days in the morning and in the afternoon for 18.75 h in total always including the onset and ending of the diurnal activity period. The nest was excavated on 5 April using a combination of two reading glasses that provided a sufficient magnification. In the field, nest dimensions were measured using a strip of millimetre paper (accuracy 1 mm). The cells with their content were separately stored in Eppendorf vials in a freezer until they were finally investigated under a Wild M3 stereomicroscope (maximum magnification times 60) on 6 August 2015. Micro-photos were taken with a Leica IC 80 HD camera mounted on a Leica MS 5 stereomicroscope (magnification times 80).
Quartinia canariensis inhabited semidesertic areas on sandy ground sparsely covered by halophytic vegetation with scattered shrubs of Salsola divaricata Masson ex Link in Buch (Amaranthaceae), Launaea arborescens (Batt.) Murb. (Asteraceae) and Zygophyllum fontanesii Webb & Berthel. (Zygophyllaceae) (Fig.
1 Habitat of Quartinia canariensis at locality III Istmo de la Pared, 0.5 km northwest of Costa Calma, Fuerteventura 2 Female of Quartinia canariensis resting on the ground near plant of Frankenia 3 Male of Quartinia canariensis standing on the ground close to Frankenia dwarf shrub 4 Nesting site at locality I Montaña Pelada. 5 Male resting in partly sand filled shell of the snail Theba geminata.
During random searching for flower visitation by Q. canariensis 15 sightings of flower visiting females, 2 sightings of flower visiting males as well as the sighting of 1 individual of unidentified sex of Q. canariensis were exclusively recorded at Frankenia laevis. Likewise, sightings of 41 females, 5 males and 7 individuals of unidentified sex were recorded during point observations at flowers of F. laevis. Visits to flowers of other plant species were not observed. In addition, males and females of Quartinia canariensis were often observed standing on the ground close to Frankenia plants (Figs
During pollen uptake at flowers of Frankenia laevis the females of Quartinia canariensis stood on the petals and occasionally also on parts of the androecium or gynoecium with their hind and mid legs, while the head was orientated towards the anthers (Fig.
Flower visiting behaviour of Quartinia canariensis at flowers of Frankenia laevis: 6 Female feeding on pollen directly from the anthers 7 Female brushing pollen from her frons with alternating movements of her fore legs 8 Nectar uptake by female: a Proboscis protruded into corolla b Head and proboscis pushed deeper down into corolla tube 9 Male feeding on pollen directly from the anthers 10 Male with protruded proboscis taking up nectar (np = nectariferous pocket).
The behaviour of the males during pollen gathering (Fig.
The pollen composition of the brood cell provisions of Quartinia canariensis is summarized in Table
Details of the brood cells of nest No.1 of Quartinia canariensis investigated on 5 April, 2015 at locality I.
Cell No. | Orientation to the north (°) | Depth below ground surface (mm) | Condition | Content | Pollen composition |
---|---|---|---|---|---|
1 | 250 | 20 | ? | pollen loaf [egg or larva probably artificially lost, as cell had been damaged during excavation] | Frankenia >99% |
2 | 340 | 20 | ? | egg at distal end of cell beyond pollen loaf | Frankenia 95%, Polycarpaea 5% |
3 | 70/250 | 20 | sealed | large larva, pollen loaf | Frankenia >99% |
4 | 110/290 | 15 | sealed | small larva, pollen loaf | Frankenia 80%, Polycarpaea 20% |
5 | 70/250 | 23 | sealed | large larva, small remnants of pollen loaf | Frankenia >99% |
Males of Quartinia canariensis frequently stood on the ground close to flowering patches of Frankenia laevis (Fig.
The behavioural sequence during copulation was subdivided into three phases: 1. initiation, 2. insertion, 3. separation. Initiation always started by a flying male pouncing on a female that was visiting flowers or resting on the ground. The male always alighted on the back of the female and managed to hold on and to orientate his longitudinal axis parallel to hers. Finally, the head of the male was situated above the mesonotum of the female, level with her tegulae. His antennae were orientated straight anterior-downwards. The fore legs of the male were placed on the female’s tegulae or on adjacent parts of her mesonotum or metanotum and his mid and hind legs were wrapped around her wings and her metasoma (Figs
Initiation phase of copulation of Quartinia canariensis: 11a Calm position of a male on the back of a female after mounting b Male trying to accomplish insertion of his genitalia while the female raises her fore legs and bends her metasoma anterior-ventrad. c Female raising hind leg against the male 12 Male vibrating vigorously with his wings during “grappling” between the partners while the female is bending her metasoma strongly anterior-ventrad 13 Female is raising her mid and hind legs against the male and bending her metasoma anterior-ventrad 14 Second male holding on to mounting male of a pair after alighting on his back.
Behaviour of the nest owning female of Quartinia canariensis at nest No. 1: 16 Female “lining” of the turret rim with the mouthparts (mandible spread, labrum erected, part of maxilla visible) at the onset of activity in the morning 17 Female backing out of the shaft during nest excavation, carrying a large sand grain behind the mandibles 18 Female turning around outside of the nest prior to the start of flight activity in the morning: a Female “lining” the turret wall from the inside b Female turning quickly around after she has left the turret head first c Tip of the metasoma of the female is visible in the nest entrance after re-entering the nest head first 19 Female turning around outside of the nest at the end of activity in the afternoon: a Female situated above the turret after she has backed out of the nest and moved forward again b Female re-entering the nest backwards.
Behavioural sequence of the copulation of Quartinia canariensis (reconstructed from a series of 16 photos).
Time (s) | Behaviour | Figure |
---|---|---|
0 | Initiation: ♂ pounced on ♀ on Frankenia flower. ♂ alighted on back of ♀; body axis of ♂ parallel to hers; head of ♂ situated above mesonotum of ♀ level with her tegulae; ♂ directed antennae straight anterior-downwards, so that ventral side of distal end of antennal club came into contact with posterior parts of pronotum and anterior parts of mesonotum of ♀; fore legs of ♂ placed laterally on scutellum and wing base of ♀, mid and hind legs of ♂ wrapped around wings and metasoma of ♀. ♀ remained passive, with antennae orientated 45° to longitudinal axis. | 15a |
32 | ♂ moved slightly backwards and shifted antennae laterally-outwards (transverse to body axis); metasoma of ♂ was turned to the right side of ♀ and was curved anterio-ventrad; ♂ genital protruded. ♀ shifted antennae laterally-outwards and opened genital chamber. | 15b |
37 | Insertion: ♂ moved antennae to a position 45° to longitudinal axis and inserted his genital in genital chamber of ♀. | 15c |
43 | ♂ changed his position on ♀ very rapidly. After that, body of ♂ curved around posterior end of ♀ metasoma, only connected to ♀ by genitalia. Head of ♂ protruded and erect, mandibles spread, antennae stretched straight forward; ♂ fore legs stretched forward, mid and hind legs bent but raised without contact to body of ♀. Wings of ♀ drooping obliquely downwards. | 15d |
52 | Head of ♂ directed downwards, mandibles of ♂ open, touching posterior margin of metasomal tergum III of ♀, antennae of ♂ directed obliquely downwards; fore, mid and hind legs of ♂ pressed against distal end of ♀ metasoma. | 15e |
58 | Head of ♂ protruded and erect, mandibles spread, antennae stretched straight forward; ♂ fore legs stretched forward, mid and hind legs bent but raised without contact with body of ♀. | as in 15d |
79 | Head of ♂ directed downwards, mandibles of ♂ open touching posterior margin of metasomal tergum III of ♀, antennae of ♂ directed obliquely downwards; fore, mid and hind legs of ♂ attached to distal end of ♀ metasoma. | as in 15 e |
93 | Head of ♂ protruded and erect, mandibles spread, antennae stretched straight forward; ♂ fore legs stretched forward, mid and hind legs bent but raised without contact with body of ♀. | as in 15d |
144 | Head of ♂ directed downwards, mandibles of ♂ open, touching posterior margin of metasomal tergum III of ♀, antennae of ♂ directed obliquely downwards; fore, mid and hind legs of ♂ pressed against distal end of ♀ metasoma. | as in 15e |
152 | Separation: ♂ genital removed from ♀ genital chamber. ♀ shifted antennae to a position 45° to longitudinal axis and moved slowly forward. ♂ dismounted ♀ postero-laterally. ♂ and ♀ flew off. | 15f |
Male-male interactions were only rarely observed: Twice a patrolling male was observed approaching and pouncing on a male that was on the ground. On both occasions this led to a short interaction before both males flew off. Furthermore, there are two observations of a second male alighting on a male that had already mounted a female and was holding on to her back (Fig.
Nest structure: The nest consisted of a subterranean burrow. The entrance was surmounted by a short cylindrical turret measuring about 2.5 mm in diameter and 1.5 mm in height (Figs
Nest No. 1 of Quartinia canariensis: 20 Close vicinity of the nest with a small fly (Diptera, Cyclorrhapha, Fam. indet.) probably inspecting the nest entrance 21 Female re-entering nest head first after backing out of the shaft prior to onset of flight activity in the morning 22 Nest entrance with dull whitish inner silken lining 23 Excavated burrow on 5 April, 2015 with turret removed and the shaft opened in the upper part to show the inner wall. The content of the opened brood cell No.2 is visible at the lower end of the shaft 24 Silk threads on and between sand grains from the wall of the nest.
The walls of the turret, the shaft and the brood cells consisted of sand grains (Figs
Brood cell content: The content of the brood cells is summarized in Table
Behaviour at the nest: During nest excavation the female backed out of the shaft carrying a load of small sand grains or a single larger particle with her mouthparts (Fig.
In the morning, at the start of diurnal activity, the female always appeared head first in the nest entrance continually treating the inner surface of the shaft and the turret with the mouthparts. These were moved slowly in close contact over the surface (Figs
The diurnal activity of the female from nest No. 1 lasted for 4.8 to 6.0 h (median 5.6 h; n = 5) starting in the morning from 10h17 to 11h24 (median 10h53; n = 5) and finishing in the afternoon from 15h48 to 17h32 (median 16h22; n = 5). The female always spent the night within her burrow. The activity of males and females at flowers of Frankenia declined noticeably early in the afternoon and was mainly over around 15h00. On a single occasion at the end of the diurnal flight period, a male was observed to enter head first into an empty snail shell of Theba geminata (Mousson). The shell was approximately 10 cm away from a flowering patch of Frankenia. Within the shell the male first performed some movements then he rolled up his metasoma and remained motionless in a resting position (Fig.
On a single occasion a small fly (Diptera, Cyclorrhapha, family indet.) was resting on the ground close to the turret, probably inspecting the nest entrance (Fig.
At all sites the only flowers that were observed to be visited by females and males of Quartinia canariensis were those of Frankenia laevis, indicating a close association with this plant that was used as a source of pollen and nectar. This is in congruence with the study by
During their visits to flowers of Frankenia laevis males and females of Quartinia canariensis usually came into contact with the anthers and stigmas of the flowers and there were often pollen grains on the exoskeleton of the wasps. Moreover, they showed a high flower fidelity to this plant. Therefore Quartinia canariensis probably acts as a pollinator of Frankenia laevis.
Males of Quartinia canariensis searched for females at flowers of Frankenia laevis, which was the main pollen source of the females. This is in congruence with mate searching in Afrotropical species of Quartinia in which the males also frequently search for females at floral resources (
Females of Quartinia canariensis avoided contact with patrolling males at flowers and offered long lasting and predominantly successful resistance against insertion by males during the initiation phase. Since the females obviously bear considerable costs to withstand the male mating efforts, it can be assumed that insertion of the male genitalia is associated with high costs for the females of Quartinia canariensis, at least during the investigated period of the flight season. The nature of these costs is still unknown.
The copulation of a Quartinia species has not been described before in detail. Compared to photos of mating pairs of Afrotropical species of Quartinia in
Quartinia canariensis was found nesting in friable sandy soil close to its main forage plant. This is very similar to the nest situation recorded for two Afrotropical species, Quartinia vagepunctata (
As in all other Afrotropical Quartinia species for which nesting is known (
The short vertical turret surmounting the nest entrance of Quartinia canariensis is of the same form as in the Afrotropical Quartinia species (
The nest of Quartinia canariensis consisted of a subterranean burrow terminated by a cell, which is principally similar to the nest architecture of Quartinia vagepunctata (
The behaviour of a Quartinia species during nest excavation has been observed for the first time. The female of Q. canariensis backed out of the shaft carrying soil particles with her mouthparts, which is similar to the behaviour of all other primarily ground nesting Masarine wasps, i.e. Priscomasaris (
The position of the egg at the distal end of the brood cell beyond the pollen loaf indicates that egg deposition precedes pollen provisioning in Quartinia canariensis. This is a plesiomorphic behavioural trait adopted from the ground pattern of the Masarinae and even the Vespidae (
In 1990 Gusenleitner established “Chrysis atracomitata LINS.” as a cleptoparasite of Quartinia canariensis. Obviously, this is a misspelling of the name Chrysis atrocomitataLinsenmaier, 1993 and the spelling has already been corrected in this sense by
Sarah Gess made very helpful comments on the manuscript and kindly corrected our English. Jim Carpenter and Robert Paxton reviewed and improved the manuscript. Annette Rosenbauer identified the collected plants. Ashley Kirk-Spriggs helped with the identification of the fly in Fig.