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Theses and Dissertations Theses and Dissertations
5-1-2009
Flagellar sensilla of females of selected species of Vespidae (Hymenoptera)
Andrew Oliver McCaskill
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FLAGELLAR SENSILLA OF FEMALES OF SELECTED SPECIES
OF VESPIDAE (HYMENOPTERA)
By
Andrew Oliver McCaskill
A Thesis Submitted to the Faculty of Mississippi State University in Partial Fulfillment of the Requirements for the Degree of Masters of Science in Entomology in the Department of Entomology and Plant Pathology
Mississippi State, Mississippi
May 2009 FLAGELLAR SENSILLA OF FEMALES OF SELECTED SPECIES
OF VESPIDAE (HYMENOPTERA)
By
Andrew Oliver McCaskill
Approved:
______Gerald T. Baker Professor of Entomology (Director of Thesis)
______Richard L. Brown Professor of Entomology (Committee Member)
______Peter W. K. Ma Associate Professor of Entomology (Committee Member)
______Clarence Collison, Professor and Head, Department of Entomology and Plant Pathology & Graduate Coordinator
______Melissa J. Mixon Interim Dean of College of Agriculture and Life Sciences Name: Andrew Oliver McCaskill Date of Degree: May 2, 2009
Institution: Mississippi State University
Major Field: Entomology
Major Professor: Dr. Gerald Baker
Title: FLAGELLAR SENSILLA OF FEMALES OF SELECTED SPECIES OF VESPIDAE (HYMENOPTERA)
Pages in Study:
Candidate for Degree of Master of Science
Little information regarding the flagellar sensilla of the Vespidae, particularly
Polistes, exists in the literature. A variety of social behaviors in vespids, such as alarm pheromones and pheromone response, involve intraspecific communication. Therefore, an understanding of sensory receptors in social wasps would benefit further behavioral and neurological studies of wasps. The flagellar sensilla of female paper wasps Polistes dorsalis, Polistes metricus, and the solitary wasp Monobia quadridens were examined by scanning and transmission electron microscopy. A number of olfactory receptors, contact chemoreceptors, and mechanoreceptors were named and described. Possible functions of these receptors were proposed based on morphological and ultrastructural data for each type of sensillum. Counts of several sensilla were made over the whole of five flagellae of each species and results were compared among species, flagellomeres, medial and lateral surfaces.
DEDICATION
I would like to dedicate this research to my wife, Miranda, and to my parents, Dan and Ann McCaskill.
ii
ACKNOWLEDGEMENTS
The author expresses sincere gratitude to those who have given their assistance in this research, and without whom this thesis would not have been possible. I would like to thank Dr. Gerry Baker for his guidance and expertise throughout my graduate program. I would also like to thank my thesis committee, Dr. Richard Brown, Dr. Peter Ma, and Dr.
Clarence Collison, for their assistance with my research. I would like to thank Dr. Jan
DuBein for assistance with the statistical analyses. Finally, I would like to thank Bill
Monroe, Amanda Lawrence, and Richard Kuklinski of MSU EM Center for their assistance with my electron microscopy work.
iii
TABLE OF CONTENTS
DEDICATION ...... ii
ACKNOWLEDGEMENTS ...... iii
LIST OF TABLES ...... v
LIST OF FIGURES ...... vii
LIST OF ABBREVIATIONS ...... ix
CHAPTER
I. INTRODUCTION ...... 1
II. MATERIALS AND METHODS ...... 6
SEM Methods ...... 6 TEM Methods ...... 7 Statistical Methods ...... 8
III. RESULTS ...... 9
IV. DISCUSSION ...... 18
V. CONCLUSION ...... 24
VI. TABLES ...... 25
VII. FIGURES ...... 34
REFERENCES CITED ...... 51
iv
LIST OF TABLES
1 Total Sensilla Basiconica A by Species ...... 25
2 Sensilla Basiconica A on Polistes metricus by Surface ...... 25
3 Sensilla Basiconica A on Polistes dorsalis by Surface ...... 25
4 Sensilla Basiconica A on Monobia quadridens by Surface ...... 26
5 Sensilla Basiconica A on Polistes metricus by Flagellomere ...... 26
6 Sensilla Basiconica A on Polistes dorsalis by Flagellomere ...... 26
7 Sensilla Basiconica A on Monobia quadridens by Flagellomere ...... 27
8 Total Plate Organs by Species ...... 27
9 Plate Organs on Polistes metricus by Surface ...... 27
10 Plate Organs on Polistes dorsalis by Surface ...... 27
11 Plate Organs on Monobia quadridens by Surface ...... 28
12 Plate Organs on Polistes metricus by Flagellomere ...... 28
13 Plate Organs on Polistes dorsalis by Flagellomere ...... 28
14 Plate Organs on Monobia quadridens by Flagellomere ...... 29
15 Total Sensilla Campaniformia by Species ...... 29
16 Sensilla Campaniformia on Polistes metricus by Surface ...... 29
17 Sensilla Campaniformia on Polistes dorsalis by Surface ...... 29
18 Sensilla Campaniformia on Monobia quadridens by Surface ...... 30
19 Sensilla Campaniformia on Polistes metricus by Flagellomere ...... 30
v
20 Sensilla Campaniformia on Polistes dorsalis by Flagellomere ...... 30
21 Sensilla Campaniformia on Monobia quadridens by Flagellomere ...... 31
22 Total Pit Organs by Species ...... 31
23 Pit Organs on Polistes metricus by Surface ...... 31
24 Pit Organs on Polistes dorsalis by Surface ...... 31
25 Pit Organs on Monobia quadridens by Surface ...... 32
26 Pit Organs on Polistes metricus by Flagellomere ...... 32
27 Pit Organs on Polistes dorsalis by Flagellomere ...... 32
28 Pit Organs on Monobia quadridens by Flagellomere ...... 33
vi
LIST OF FIGURES
1 Monobia quadridens antenna...... 34
2 Polistes dorsalis antenna...... 34
3 Polistes metricus antenna...... 34
4 Significant differences in s. basiconica A by flagellomere. M. quadridens...... 35
5 Significant differences in s. basiconica A by flagellomere. P. dorsalis...... 35
6 Significant differences in s. basiconica A by flagellomere. P. metricus...... 35
7 Significant differences in plate organs by flagellomere. M. quadridens...... 36
8 Significant differences in plate organs by flagellomere. P. dorsalis...... 36
9 Significant differences in plate organs by flagellomere. P. metricus...... 36
10 Significant differences in pit organs by flagellomere. M. quadridens...... 37
11 Significant differences in pit organs by flagellomere. P. dorsalis...... 37
12 Significant differences in pit organs by flagellomere. P. metricus...... 37
13 Significant differences in s. campaniformia by flagellomere. M. quadridens...... 38
14 Significant differences in s. campaniformia by flagellomere. P. dorsalis...... 38
15 Significant differences in s. campaniformia by flagellomere. P. metricus...... 38
16 Plate Organs...... 39
17 Sensilla Basiconica A ...... 40
18 Sensilla Basiconica B...... 41
vii
19 Sensilla Campaniformia...... 42
20 Pit Organs...... 43
21 Sensilla Trichodea A ...... 44
22 Sensilla Trichodea B...... 45
23 Sensilla Chaetica ...... 46
24 Sensillum Spatulatum and Setae of M. quadridens ...... 47
25 Setae of P. dorsalis ...... 48
26 Setae of P. metricus ...... 49
27 Sections of Sensilla of P. dorsalis ...... 50
viii LIST OF ABBREVIATIONS
A Setae A
B Setae B
Ba Sensilla Basiconica A
Bb Sensilla Basiconica B
C Setae C
Ca Sensilla Campaniformia
Ch Sensilla Chaetica
D Setae D
De Dendrites
L Lumen
Pi Pit Organs
Pl Plate Organs
Sp Sensilla Spatulata
Ta Sensilla Trichodea A
Tb Sensilla Trichodea B
↑ Wall Pores
ix
CHAPTER I
INTRODUCTION
The family Vespidae is a large group of solitary and eusocial wasps with 325 species in 5 subfamilies in North America (Triplehorn and Johnson 2005). Two of the largest subfamilies are Polistinae and Eumeninae. Subfamily Polistinae is a large and diverse group of primitively eusocial wasps with around 800 described species in 29 genera worldwide (Carpenter 1991, Triplehorn and Johnson 2005) of which the genus
Polistes includes with 206 described species (Akre 1982). In North America, there are
325 species of Vespidae, with 17 of those occurring in Polistes (Triplehorn and Johnson
2005). Subfamily Eumeninae is relatively large, with 206 species in North America
(Triplehorn and Johnson 2005) and the genus Monobia contains one North American species (Buck et al. 2008).
The wasps of Polistes are of interest due to their social behavior (Reeve 1991). In
Polistes there is not a clear morphological distinction between castes (Eberhard 1969).
Queens and workers differ little in size when compared to some other social
Hymenopterans, specifically bees and some of the Vespa (Evans 1958, Andrewes 1969).
One primary difference between Polistes castes is that workers do not have fully developed ovaries while the queen does (Andrewes 1969).
1
Foundress wasps overwinter and emerge in the spring to build nests (Spradbery
1973). Foundresses forage for materials for nest construction, hunt for prey, and provide care for the first few immatures until they develop to adults, at which point a division of labor begins to appear (Spradbery 1973). New workers assume foraging for food and nest materials, hunting for food, and nest building activities while the foundress gradually stops hunting and focuses on nest construction and egg laying (Spradbery 1973). As the season progresses, males and reproductive females are produced (Reeve 1991). Mating occurs upon their maturation and the colony cycle ends with the death of the workers and males while the fertilized reproductive females overwinter (Andrewes 1969).
Workers forage for plant fibers and water for nest construction, nectar for feeding the adults of the nest, and prey for feeding the immatures of the nest (Richter 2000).
Vespid wasps prey upon a number of different insects, spiders, and occasionally vertebrate carrion (Richter 2000) which, once captured, they masticate and feed to the immature larvae via trophyllaxis (Spradbery 1973).
A considerable amount of literature on Vespidae deals with various chemicals that are involved with behavior. Volatiles such as pheromones play an important role in the behavior of wasps. These include chemical trails, alarm pheromones, pheromones related to mating and courtship behavior, cuticular lipids used in nestmate recognition, local enhancement, and the location of prey. Cavity-nesting Vespidae such as Vespa crabro and Vespula vulgaris use chemical trails to locate nests because the cavity entrance is often spatially separated from the nest entrance (Steinmetz et al. 2002). Many vespids also use an alarm pheromone (Landolt et al. 1997) to initiate a response in nestmates
2
(Reed and Landolt 2000). Alarm pheromones are present in the venom glands of many vespids, including Polistes dominulus (Bruschini et al. 2006), Vespula maculifrons
(Landolt et al. 1995), Vespula squamosa (Landolt and Heath 1987), and in the head of at least one vespid, Vespula squamosa (Landolt et al. 1999). In V. crabro and P. dominulus males, secretory cells and release structures are found in some flagellomeres (Romani et al. 2005). Cuticular lipids are necessary for nestmate recognition (Howard 1993). Dani et al. (2001) found that methyl branched alkanes and alkenes were used by P. dominulus to recognize nestmates. Cuticular lipids are species- and colony-specific (Espelie and
Hermann 1988), and the hydrocarbons found on the nests and are necessary for nest recognition in social wasps (Espelie and Hermann 1990). Environmental factors play a role in cuticular hydrocarbon profiles in P. metricus (Singer and Espelie 1997) and may play a role in P. fuscatus (Gamboa et al. 1996) and P. dominulus (Dani et al. 2001). It has been suggested that the yellow jacket Vespula germanica uses volatiles as olfactory cues coupled with visual cues in foraging for food and other resources (D’Adamo et al.
2003). Richter (2000), in a review of the literature, suggests that olfactory cues play at least some part in locating prey items in social wasps.
Antennation, the touching of a surface with the antennae (Romani et al. 2005), plays an important role in many social behaviors observed in wasps. Antennation occurs during courtship and mating. The male wasp hooks his antennae around the antennae of the female and rubs his up and down the length of hers while mating (Romani et al.
2005). The secretory cells reported by Romani et al. (2005) may play a role in this mating behavior. Antennation also occurs during dominance interactions in Polistes
3 gallicus (Pardi 1948); during trophyllaxis in Polistes fuscatus (Eberhard 1969); and while building new cells in a nest (Eberhard 1969).
Few publications regarding flagellar sensilla in Vespidae are present. Callahan
(1975) presents several SEM micrographs of the flagellar sensilla of the vespid Polistes metricus, as well as numerous micrographs of the flagellar sensilla of many other species of insects in several different orders. Spradbery (1973) also provides several SEM micrographs of the flagellar sensilla of Vespula rufa and V. crabro.
There are several investigations of flagellar sensilla in the closely related family
Pompilidae. Micrographs and descriptions of the shape and metrics of each type of sensillum, possible functions, and comparisons with sensilla found in other Hymenoptera have been presented for Anopilius tenebrous (Alm and Kurczewski 1982), Evagetes parvus (Lane et al. 1988), and Poecilagenia sculpturata (Shimizu 2000). Alm and
Kurczewski (1982) propose functions for the sensilla found during their study of A. tenebrous by comparing them to structurally similar sensilla found in other Hymenoptera.
Shimizu (2000) correlates the function of sensilla on the antennae of P. sculpturata to cleptoparasitic behavior. The female of this species applies the ventral portion of her flagella, which are more densely covered in sensilla basiconica A, to a substrate when searching for hosts or stored spiders, therefore these sensilla are of significant use in searching behavior (Shimizu 2000).
Monobia quadridens is a solitary wasp in which the females nests in wood cavities made by other insects, such as carpenter bees (Krombien 1967). M. quadridens females provision their nests with caterpillars and build partitions between cells with mud
4 or sand (Krombien 1967). During mating, males will seek out locations where females are nesting and wait for them, performing dances prior to mating (Krombien 1967).
5
CHAPTER II
MATERIALS AND METHODS
Specimens of adult females were collected in the field, either individually
(Monobia) or along with their nests (Polistes), or were reared from collected nests
(Polistes). The antennae of Polistes dorsalis females were processed for SEM and TEM.
The antennae of Polistes metricus and Monobia quadridens females were processed for
SEM only. The antennae of a male of P. dorsalis and of P. metricus were also processed for SEM. The SEM and TEM processing procedures follow the basic protocols from
Bozzola and Russell (1999) with some modifications and are described below.
SEM Methods
Antennae were cut from living specimens were fixed overnight in a triple aldehyde solution containing 5% glutaraldehyde, 4% paraformaldehyde, and 2% acrolein in 0.1 M phosphate buffer, pH 7.2, at 4° C. Antennae were rinsed in buffer three times,
20 minutes each, the antennae were post-fixed in 2% OsO4, with the same buffer, for three hours on ice and one hour at room temperature. Antennae were rinsed three times in distilled water, 10 minutes each, and dehydrated in a graded series of ethanol with two
10 minute changes of 35%, 50, and 75% ethanol, two 15 minute changes of 95% ethanol, and four 15 minute changes of 100% ethanol. After being dehydrated, antennae were in
6
HMDS for two 10 minute rinses and allowed to air dry. The antennae were mounted on aluminum stubs with double sided carbon tape and silver paste adhesive. The antennae were sputter coated for ~1 minute with gold palladium. The stubs were examined with a
JEOL-JSM 6500F at 5 kV, and the images were digitally recorded. The digitally recorded micrographs were used to determine the types of flagellar sensilla present and to make counts of several sensilla types.
TEM Methods
Antennae cut from living specimens were cut further into smaller pieces and fixed overnight in a triple aldehyde solution containing 5% glutaraldehyde, 4% paraformaldehyde, and 2% acrolein in 0.1 M phosphate buffer, pH 7.2, at 4° C. After three buffer rinses, 20 minutes each, the antennae were post-fixed in 2% OsO4, with the same buffer, for three hours on ice and one hour at room temperature. After post- fixation, the antennae were rinsed in distilled water, with three 20 minute changes, and stained en bloc overnight in uranyl acetate. After being rinsed in distilled water, three 20 minute changes, the antennae were dehydrated in a graded series of ethanol with two 10 minute changes of 35%, 50, and 75% ethanol, two 15 minute changes of 95% ethanol, and with four 15 minute changes of 100% ethanol. The antennae then were placed in a transitional fluid, 100% acetone, for 30 minutes, before they were infiltrated with Spurr’s low viscosity embedding medium. A graded series of acetone and Spurr’s medium was used over a 48 hour period. The infiltrated specimens were placed in molds and put in an oven at 70°C overnight to polymerize. The tissue blocks were trimmed and thick
7 sections (1μ) were cut, slide mounted, and stained with toluidine blue to locate the areas of interest. Thin sections (65-70 nm) were cut and were placed on formvar coated copper grids, stained with uranyl acetate (20 minutes) and lead citrate (10 minutes). The stained sections were viewed with a JEOL JEM-100CXII TEM scope at 60-80kV, and micrographs were recorded on Kodak plate film.
Statistical Methods
The number and distribution of sensilla were recorded for the selected species.
Monobia was chosen as an out-group for comparison purposes. Sensilla basiconica A, plate organs, sensilla campaniformia, and pit organs were chosen for making counts for 5 flagella in each species. The flagellum consists of ten flagellomeres, or subsegments, and two surfaces, medial and lateral. Means were compared using Student-Newman-Keuls at the 5% significance level.
8
CHAPTER III
RESULTS
The antennae of female vespids have two basal segments, the scape and pedicel, and ten additional flagellomeres that are covered with setae and sensilla (Figures 1-3).
The 10 flagellomeres are densely covered in setae, non-innervated hairs (Lane et al.
1988), and various sensilla, innervated structures with a sensory function. In this study, the flagellomeres are numbered from 1 to 10 beginning proximally. The antennae may be divided into two distinct surfaces based on morphological characteristics and these distinct surfaces correspond to the position of the antennae. The two surfaces will be referred to as the medial and lateral surfaces, based on the position of the antennae when held at rest in front of and to the side of the head. The medial surface faces towards the head, and the lateral surface faces away from the head when held at rest.
The nomenclature of sensilla is taken from Shimizu (2000), with some modifications. Generally, all sensilla either project out perpendicular to the long axis of the antenna, or at varying acute angles with their apices pointing towards the distal end of the flagellum. All sensilla found were given a designation and described. Though unlikely, sensilla other than those encountered may be present on the antennae.
Plate organs (Figures 16, 27B, and 27C) are long smooth multiporous structures that are oriented longitudinally on the flagellum. The pore plate is surrounded by a
9 fissure and the distal end comes to a pointed keel while the proximal end is rounded. The surface is covered by rows of fine pores set apart at regular intervals. TEM investigation shows that, in P. dorsalis, the sensillum is thin-walled, and multiple branching dendrites extend through the lumen to the multiporous cuticle (Figures 27B and 27C). The mean length of the plate organ is 17.14 μm (12.29 – 21.04 μm) in P. metricus, 19.35 μm (16.97
– 21.38 μm) in P. dorsalis, and 15.69 μm (15.39 – 16.05 μm) in M. quadridens. The mean width of the plate organ is 3.88 μm (2.81 – 4.50 μm) in P. metricus, 3.79 μm (2.72
– 4.50 μm) in P. dorsalis, and 4.59 μm (4.19 – 5.18 μm) in M quadridens. The plate organs are numerous and cover almost the entire surface of each flagellomere, except for the apex of the 10th flagellomere, the basal and apical margin of the 2nd through 10th flagellomere, and a portion of the 1st flagellomere. On the antennae of Monobia, they are absent from a portion of the 2nd flagellomere. Plate organs are found on all other species investigated and on all flagellomeres.
The total number of plate organs is significantly different between all three species, with M. quadridens having significantly more and P. dorsalis having significantly fewer than P. metricus (Table 8). There are significantly more plate organs on the medial surface than on the lateral surface of the antennae of P. metricus and M. quadridens (Table 9 and 11), but there is no significant difference between surfaces in P. dorsalis (Table 10). The highest concentration of plate organs is on the medial flagellomeres, with significantly fewer plate organs on the apical and basal flagellomeres
(Table 12, 13, and 14) (Figures 7, 8, and 9).
10
Sensilla basiconica A (Figures 17 and 27A) are stout, slightly grooved longitudinally, peg-like chemosensory organs that are set into the proximal end of oval depression surrounded by a raised collar. The apex of this sensillum is often somewhat flattened or concave and occasionally with a granular texture. In P. dorsalis, the peg has a thin cuticular wall with multiple branching dendrites running up through the lumen to the multiporous apex of the sensillum (Figure 27A). The conical portion projects outward almost perpendicular to the long axis of the antennae. The mean height of the peg is 10.87 μm (8.91 – 12.04 μm) in P. metricus, 11.98 μm (10.50 – 13.68 μm) in P. dorsalis, and 6.39 μm (5.54 – 6.82 μm) in M. quadridens; the mean width of the peg at its base is 7.34 μm (5.88 – 8.31 μm) in P. metricus, 7.42 μm (6.16 – 8.65 μm) in P. dorsalis, and 7.57 μm (7.15 – 8.10 μm) in M. quadridens; and the mean width of the raised collar at its widest is 11.92 μm (11.05 – 12.58 μm) in P. metricus, 11.11 μm (10.21 – 13.90 μm) in P. dorsalis, and 10.36 μm (9.52 – 11.48 μm) in M. quadridens. Sensilla basiconica A are more numerous on the medial surface than on the lateral surface, and are more common on the apical portion of the flagellum than the basal portion. They are located within the plate organ region immediately distad the sensilla basiconica B region that surrounds the base of the flagellum on flagellomeres 2 through 10. They are absent from the apex of the 10th flagellomere, and from a large portion of the 1st flagellomere. They are present in all three species, and are present on all flagellomeres.
No significant difference exists in the total number of s. basiconica A between any of the three species (Table 1). In all three species, there are significantly more s. basiconica A on the medial surface than the lateral surface (Tables 2, 3, and 4) and
11 significantly more s. basiconica A on the apical flagellomeres. The number tends to decrease towards the basal flagellomeres in all three species (Tables 5, 6, and 7) (Figures
4, 5, and 6).
Sensilla basiconica B (Figure 18) are smooth, multiporous cone-shaped organs lacking grooves and are set into a round depression. Minute pores are visible randomly over the surface of this sensillum. This sensillum is narrower and has a more pointed apex than s. basiconicum A. The conical portion of this sensillum is similar to that of s. basiconicum A in projecting outward perpendicular to the long axis of the antenna. The mean height of the sensillum is 10.25 μm (8.66 – 12.43 μm) in P. metricus, 10.30 μm
(8.44 – 11.85 μm) in P. dorsalis, and 10.75 μm (10.42 – 11.21 μm) in M. quadridens; the mean diameter of the sensillum at its base is 3.45 μm (2.78 – 4.24 μm) in P. metricus,
3.29 μm (2.81 – 3.67 μm) in P. dorsalis, and 2.32 μm (1.91 – 2.67 μm) in M. quadridens; and the mean width of the depression is 4.59 μm (4.06 – 5.24 μm) in P. metricus, 4.91
μm (4.30 – 5.40 μm) in P. dorsalis, and 4.23 μm (3.70 – 4.59 μm) in M. quadridens. The sensilla are generally found in a ring along the basal portion of the lateral surface of flagellomeres 2 through 10 on the Polistes spp. and on Monobia, though they may be located elsewhere. On the Polistes spp. the ring of s. basiconica B completely surrounds the base of flagellomeres 6 through 10, but does not completely surround the base of flagellomeres 2 through 5. On Monobia the s. basiconica B densely cover a region throughout the length of each flagellomere on the lateral surface of flagellomeres 2 through 5. They are present in all species investigated, and are present on all flagellomeres.
12
Sensilla campaniformia (Figure 19) consist of round, slightly raised granulated papillae set in the center of flat smooth round shallow indentations on the antennal surface. The mean diameter of the papilla is 1.41 μm (0.99 – 1.88 μm) in P. metricus,
1.45 μm (1.09 – 1.88 μm) in P. dorsalis, and 1.60 μm (1.48 – 1.83 μm) in M. quadridens.
The mean diameter of the depression at its widest is 9.83 μm (8.44 – 10.62 μm) in P. metricus, 9.14 μm (6.43 – 11.12 μm) in P. dorsalis, and 9.96 μm (7.58 – 11.75 μm) in M. quadridens. Although the sensilla occur on most flagellomeres of the flagellum, s. campaniformia are uncommon and can be found centrally on flagellomeres 2 through 10 in the Polistes spp., and in Monobia. They are present in all species investigated, and on most flagellomeres.
No significant difference in the total number of s. campaniformia occurs between any of the three species (Table 15). Significantly more s. campaniformia are on the lateral surface than on the medial surface of the flagellum of P. metricus (Table 16), but no significant difference in the number of s. campaniformia exists between flagellar surfaces in P. dorsalis or M. quadridens (Table 17 and 18). No significant difference in the number of s. campaniformia exists between any of the flagellomeres of P. metricus
(Table 19) (Figure 15). Significantly more s. campaniformia are present on the 10th flagellomere of P. dorsalis, but no significant difference exists between any of the other flagellomeres (Table 20) (Figure 14). Similarly, significantly more s. campaniformia are located on the 9th flagellomere of M. quadridens, but no other flagellomeres are significantly different (Table 21) (Figure 13).
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Pit organs, (sensillum coeloconicum and/or ampullaceum) (Figures 20 and 27D) appear as an opening at the center of smooth round concave indentation on the surface of the antennae. Sensilla coeloconica and sensilla ampullacea appear similar and cannot be distinguished easily by SEM alone. Rarely, it was possible to view a portion of the sensory peg within the pit with SEM. The sensory peg resembles that of sensilla coeloconica I described by Bleeker et al. (2004) in Cotesia glomerata and Cotesia rubecula (Braconidae). S. coeloconica and s. ampullacea can be distinguished in other species either by the diameter of the pit opening or by the depth of the pit. S. coeloconica have a smaller pit opening than s. ampullacea in E. parvus (Lane et al. 1988). S. coeloconica have a shallower pit than do s. ampullacea in Apis mellifera (Lacher 1964).
TEM results show that P. dorsalis possesses s. coeloconica, and the sensory peg is located within a cavity under the cuticular surface of the flagellum (Figure 27D). The mean diameter of the opening is 1.61 μm (1.15 – 1.88 μm) in P. metricus, 1.57 μm (0.94
– 2.40 μm) in P. dorsalis, and 1.80 μm (1.60 – 1.96 μm) in M. quadridens. The mean diameter of the depression at its widest is 10.68 μm (8.25 – 13.73 μm) in P. metricus,
9.64 μm (7.13 – 11.77 μm) in P. dorsalis, and 7.71 μm (7.07 – 8.75 μm) in M. quadridens. They are located in the same areas as sensilla campaniformia, and often occur in clusters in the species investigated. These are present on all three species and are found on most flagellomeres.
Significantly more pit organs are on M. quadridens than on either of the Polistes spp., which are not significantly different from each other (Table 22). Significantly more pit organs are found on the lateral surface than on the medial surface of the antennae of P.
14 metricus (Table 23), but no significant difference occurs in the number of pit organs between antennal surfaces in P. dorsalis or M. quadridens (Table 24 and 25). In all three species, significantly more pit organs are located on the apical flagellomeres, excluding the 10th, and fewer on the basal flagellomeres (Table 26, 27, and 28) (Figures 10, 11, and
12).
Sensilla trichodea A (Figure 21) are long slender sensilla that project perpendicular to the long axis of the antennae. The apex is rounded, and a pore is located at the tip of this sensillum. Scanning electron micrographs of broken s. trichodea A reveal them to be hollow. They are somewhat variable in appearance, with some having prominent longitudinal grooves or furrows, others having a relatively smooth surface, and a few having a portion of their surface grooved and the rest smooth. There may be several subtypes present, but they are difficult to distinguish from each other. The mean height of the sensillum is 17.36 μm (15.60 – 20.99 μm) in P. metricus, 15.56 μm (13.89 –
17.74 μm) in P. dorsalis, and 17.46 μm (11.05 – 22.22 μm) in M. quadridens. The mean diameter of the sensillum at its base is 2.09 μm (1.70 – 2.39 μm) in P. metricus, 1.80 μm
(1.61 – 2.08 μm) in P. dorsalis, and 2.31 μm (1.39 – 3.47 μm) in M. quadridens. They are found at intervals over the flagellum. Sensilla trichodea A occur in all three species and are present on all flagellomeres.
Sensilla trichodea B (Figures 22) are short and slender organs that project perpendicular to the long axis of the antennae. They are similar in appearance to sensilla trichodea A, but much shorter and usually have definite grooves. The mean height of the sensillum is 10.13 μm (9.30 – 10.79 μm) in P. metricus, 10.31 μm (9.31 – 11.07 μm) in
15
P. dorsalis, and 6.98 μm (6.27 – 7.42 μm) in M. quadridens. The mean diameter of the sensillum at its base is 2.35 μm (1.66 – 2.67 μm) in P. metricus, 2.20 μm (1.81 – 2.54
μm) in P. dorsalis, and 2.07 μm (1.81 – 2.52 μm) in M. quadridens. They occur in all species investigated, and are present on all flagellomeres.
Sensilla chaetica (Figure 23) are slender grooved sensilla that basally project perpendicular to the long axis of the antenna, but bend sharply so that the apical ⅔ to ¾ runs parallel to the long axis of the antenna. No pores are visible on the surface in SEM.
The apical end of this sensillum is pointed. The mean length of the sensillum is 20.06
μm (16.50 – 26.05 μm) in P. metricus, 17.58 μm (14.55 – 22.44 μm) in P. dorsalis, and
9.44 μm (8.42 – 10.65 μm) in M. quadridens. Sensilla chaetica are found at intervals along the whole flagellum. Their appearance is somewhat variable, particularly in
Monobia. They occur in all three species and are present on all flagellomeres.
Sensilla spatulata (Figure 24A) are grooved with an oval-cylindrical base and an apical projection. They project out perpendicular to the long axis of the antennae, and are present only in Monobia, occurring on all flagellomeres. The mean height of the sensillum is 9.04 μm (8.36 – 10.03 μm), mean diameter of the sensillum at its base is 2.39
μm (1.98 – 2.59 μm), and mean height of the projection 2.77 μm (2.32 – 3.30 μm).
Setae, non-innervated hairs lacking a sensory function, densely cover the surface of the flagellum in all species. Several subtypes are evident in Polistes. Setae A (Figures
24B, 25A, and 26A) are grooved with a pointed tip and occur over most of the flagellum.
Setae B (Figures 24C, 25B, and 26B) are grooved with an extended sickle-shaped tip and are common on the apical flagellomeres. Setae C (Figures 25C and 26C) are grooved,
16 curved, and occur near the distal margin of each flagellomere excluding the 10th. Setae D
(Figures 25D and 26D) are long with a pointed tip and occur on the basal two flagellomeres. Scanning electron micrographs of broken setae reveal that they are not hollow, and no pores are visible on the surface.
A brief examination of the antennae of a male P. dorsalis wasp and P. metricus revealed that the flagellomeres possess an area devoid of sensilla and setae similar to that of P. dominulus (Romani et al. 2005). Not enough specimens were collected for a thorough examination of the flagellar sensilla present on the antennae of males of both
Polistes species investigated.
17
CHAPTER IV
DISCUSSION
The functions of the various sensilla can only be determined truly through experimentation (Schneider 1964). However, morphological and ultrastructural characteristics and distributional patterns may be used to suggest possible functions
(Schneider 1964). Sensilla may function as mechanoreceptors, contact chemoreceptors
(gustatory receptors), and/or olfactory receptors. Mechanoreceptive sensilla generally lack pores (Keil 1999) and are innervated by a single dendrite (McIver 1975).
Mechanoreceptive hair sensilla may function solely as mechanoreceptors or may have an additional chemosensory function (McIver 1975). Contact chemoreceptors, or thick walled chemoreceptors, possess a terminal pore (Barbarossa et al. 1998, Zacharuk 1980,
Chapman 1998) and unbranching dendrites that terminate near the pore (Slifer 1970).
Olfactory receptors, or thin walled chemoreceptors, possess multiple wall pores (Keil
1999, Zacharuk 1980, Chapman 1998) and multiple branching dendrites that terminate near those pores (Slifer 1970).
Plate organs, also called sensilla placodea or multiporous plate sensilla (MPS) are common throughout Hymenoptera (Slifer 1970). Kaissling and Renner (1968) and
Lacher and Schneider (1963) found by experiment that the plate organs in Apis serve as olfactory receptors. Some authors suggest that plate organs may act as infrared receptors
18 in certain parasitic wasps (Borden et al. 1978a, b). The parasitic wasp Coeloides brunneri Vierick (Braconidae) has been shown to use heat to locate oviposition sites
(Richerson and Borden 1972). Richerson et al. (1972) postulate that sensilla placodea on
C. brunneri function as infrared receptors. However, the arrangement of dendrites within the sensilla placodea of C. brunneri is different from that of P. dorsalis. In C. brunneri, dendrites extend throughout the length of the sensillum (Richerson et al. 1972) whereas in P. dorsalis, the multiple branching dendrites extend through the lumen and terminate near the porous plate. Most current authors suggest that plate organs are olfactory receptors (Basibuyuk and Quicke 1999). The morphological characteristics of plate organs on the antennae of P. dorsalis suggest an olfactory function. According to Akre and Getz (1993), plate organs in Apis mellifera are well-suited for perceiving a variety of odor molecules. Plate organs in wasps could be potentially useful in finding resources
(Weseloh 1972), nestmate discrimination, as well as chemical communication. Schneider and Steinbrecht (1968) found two types of olfactory cells in the plate organs of bees, generalist and specialist. Specialist olfactory cells likely perceive pheromones.
Schneider and Steinbrecht (1968) suggest that generalist olfactory cells in the plate organs of bees perceive plant odors and fatty acids. The cuticular hydrocarbon profile of social wasps used in nestmate discrimination is similar to that of their natal nests (Espelie and Hermann 1988; 1990). Gamboa et al. (1986) found evidence that compounds in nest materials may play a role in kin recognition and nest discrimination. If plate organs perceive similar compounds in social wasps, then they may be responsible for processing the hydrocarbons used in nest and nestmate recognition. Since plate organs are likely
19 responsible for the olfaction of many different compounds for many different purposes, behavior alone may not account for the significant differences in the numbers of plate organs on the antennae of the species investigated. It is possible, however, that these differences are related to the size and surface area of the antennae. P. dorsalis, the smallest species in its genus in eastern North America (Buck et al. 2008) also has significantly fewer plate organs than the other species investigated.
Sensilla basiconica A are chemoreceptors. Morphological and ultrastructural characteristics support the assertion that sensilla basiconica A are olfactory receptors.
The sensillum is thin walled and multiporous with multiple branching dendrites. There is some uncertainty in the literature with regards to function. Shimizu (2000), investigating
Poecilagenia sculpturata, proposes an olfactory function or a gustatory (contact chemoreceptor) function for sensilla basiconica A. Alm and Kurczewski (1982) states that sensilla basiconica A (called corrugated conical sensilla) on the antennae of Anoplius tenebrosus resemble taste bristles. Lane et al. (1988) propose that s. basiconica A (called corrugated conical sensilla) are gustatory receptors in the pompilid Evagetes parvus.
Confusion may stem from the fact that the multiple pores are only located on the tip of the sensillum, similar to the terminal pore in contact chemoreceptors. The significantly higher concentration of s. basiconica A on the medial facing and on the terminal flagellomeres may also suggest a contact chemosensory function, as wasps are known to use their antennae as “feelers” and antennation occurs during a number of social interactions (i.e. dominance interactions, trophyllaxis). If s. basiconica A are olfactory receptors, then their concentration and location on the flagellum may indicate
20
“directionality” and would likely be useful in resource finding and hunting prey. Other authors, such as Lane et al. (1988) and Shimizu (2000) have suggested that s. basiconica
A are olfactory sensilla that are used in locating prey or hosts.
Sensilla basiconica B are chemoreceptors. Examination of the surface of the sensillum with SEM reveals characteristics associated with olfactory receptors, including multiple pores and thin cuticular walls seen in broken sensilla.
Sensilla trichodea A and B are contact chemoreceptors. They possess terminal pores and broken sensilla are hollow. Assuming sensilla basiconica A are olfactory receptors, then this would leave sensilla trichodea as the primary source of contact chemoreception on the antennae of the wasps investigated.
The sensilla spatulata of M. quadridens appear similar to s. trichodea, and may also be contact chemoreceptors. Lane et al. (1988) observed s. spatulata on E. parvus and suggested a chemosensory function. S. spatulata were reported in two cleptoparasitic
Pompilids, E. parvus (Lane et al. 1988) and P. sculpturata (Shimizu 2000). Behavior may explain the function of s. spatulata. S. spatulata may play a role in sensing soil. The sensilla are not located in either of the Polistes species which build nests out of plant fibers, but are found in Monobia, which builds cells from mud or sand. The two aforementioned cleptoparasitic Pompilids search for hosts and stored spiders by applying their antennae to the soil (Lane et al. 1988, Shimizu 2000). Further experiments are necessary to understand the function of this uncommon sensillum.
Sensilla chaetica are probably mechanoreceptors which lack pores and are not hollow. They seem distinct from the different setae covering the flagellum, though there
21 is not yet conclusive TEM evidence for a possible function. A similarly curved sensillum
(called s. trichodea B) although lacking grooves, occurs in Colletes cunicularis (Agren
1977) and two species of Andrena (Agren 1978) and is thought to be a mechanoreceptor
(Agren 1978). Lacher (1964) found through electrophysiological research that similar sensilla (called small thick walled hairs) in Apis mellifera have a mechanoreceptive function. However, Lane et al. (1988) found a structure similar to s. chaetica (called s. trichodea B) on E. parvus, but could not find any evidence of a mechanoreceptive function. If at some later time these structures are found to be non-innervated, then they should be considered another type of setae.
Pit organs, sensilla coeloconica and/or ampullacea, may be humidity, temperature, or CO2 receptors (Altner and Prillinger 1980). Lacher (1964) found that some pit organs in A. mellifera function as hygroreceptors and others as CO2 receptors. Sensilla campaniformia are likely mechanoreceptors. Dietz and Humphreys (1971) suggest that s. campaniformia in honey bees are related to s. coeloconica/ampullaceal and have a similar function. Other authors, however, posited a mechanoreceptive function. Lane et al.
(1988) found that s. campaniformia in E. parvus were proprioceptors.
Evidence from SEM and TEM examination suggests that the several types of setae on the flagellum are not innervated. They lack pores, are not hollow, do not appear to have any underlying structures associated with mechanoreceptive sensilla, and they closely resemble non-innervated hairs found in other Hymenoptera, including the
Apoidea (Wcislo 1995) and on the antennae of Pompilid Evagetes parvus (Lane et al.
1988). If at some point more conclusive evidence is found that one or more of the types
22 of setae present are innervated, then those setae should be considered sensilla chaetica.
Spradbery (1973) indicates that setae found on the body and appendages of social wasps function as mechanoreceptors or are used in grooming depending on their location.
Sumana and Starks (2004) reported that Polistes dominulus wasps groomed themselves with their legs, wings, and mouthparts, but did not mention that the antennae were used in grooming. If any of the setae have a mechanoreceptive function, they may play some role in antennation during the construction and shaping of nest cells.
23
CHAPTER V
CONCLUSION
Functions proposed for the sensilla above vary in the level of certainty. Some sensilla, such as plate organs, have been investigated in detail in a number of insects and there is evidence for an olfactory function. Other receptors, such as sensilla basiconica B and sensilla spatulata, are less common and not well represented in literature. Though no experiments were performed to determine function in this study, the ultrastructural details of the several sensilla investigated do suggest possible functions. A thorough investigation of the males of these three species should be undertaken, as similarities and differences in the type, number, and distribution of flagellar sensilla between sexes may provide further evidence for the function of sensilla. Additionally, the flagella of males should be studied to determine if they have secretory cells similar to those of Polistes dominulus (Romani et al. 2005), and what potential roles their secretions play in social behaviors such as mating. Future studies should work to experimentally determine function and relate that information to morphology and ultrastructure as well as insect behavior.
24
CHAPTER VI
TABLES
Table 1: Total Sensilla Basiconica A by Species
Species N Mean Monobia quadridens 5 490.6 a Polistes metricus 5 601.0 a Polistes dorsalis 5 518.4 a Means followed by same letter are not significantly different
Table 2: Sensilla Basiconica A on Polistes metricus by Surface
Surface N Mean Medial 5 478.8 a Lateral 5 122.2 b Means followed by same letter are not significantly different
Table 3: Sensilla Basiconica A on Polistes dorsalis by Surface
Surface N Mean Medial 5 409.2 a Lateral 5 109.2 b Means followed by same letter are not significantly different
25
Table 4: Sensilla Basiconica A on Monobia quadridens by Surface
Surface N Mean Medial 5 365.8 a Lateral 5 124.8 b Means followed by same letter are not significantly different
Table 5: Sensilla Basiconica A on Polistes metricus by Flagellomere
Flagellomere N Mean 10 5 95.4 a 9 5 79.6 b 8 5 77.8 b 7 5 75.4 b 6 5 62.4 c 5 5 54.2 cd 4 5 45.6 de 3 5 42.2 ef 2 5 37.8 ef 1 5 30.6 f Means followed by same letter are not significantly different
Table 6: Sensilla Basiconica A on Polistes dorsalis by Flagellomere
Flagellomere N Mean 10 5 74.0 a 9 5 68.8 ab 8 5 68.6 ab 7 5 60.4 b 6 5 57.0 b 5 5 46.4 c 4 5 39.2 cd 3 5 38.8 cd 2 5 36.4 cd 1 5 28.8 d Means followed by same letter are not significantly different
26
Table 7: Sensilla Basiconica A on Monobia quadridens by Flagellomere
Flagellomere N Mean 10 5 124.8 a 9 5 131.4 a 8 5 88.0 b 7 5 59.8 bc 6 5 30.8 cd 5 5 21.6 cd 4 5 14.2 d 3 5 9.0 d 2 5 7.0 d 1 5 4.0 d Means followed by same letter are not significantly different
Table 8: Total Plate Organs by Species
Species N Mean Monobia quadridens 5 4634.2 a Polistes metricus 5 3700.2 b Polistes dorsalis 5 2481.2 c Means followed by same letter are not significantly different
Table 9: Plate Organs on Polistes metricus by Surface
Surface N Mean Medial 5 2052.8 a Lateral 5 1647.4 b Means followed by same letter are not significantly different
Table 10: Plate Organs on Polistes dorsalis by Surface
Surface N Mean Medial 5 1270.0 a Lateral 5 1211.2 a Means followed by same letter are not significantly different
27
Table 11: Plate Organs on Monobia quadridens by Surface
Surface N Mean Medial 5 2677.8 a Lateral 5 1956.4 b Means followed by same letter are not significantly different
Table 12: Plate Organs on Polistes metricus by Flagellomere
Flagellomere N Mean 10 5 290.4 b 9 5 328.6 ab 8 5 347.2 ab 7 5 354.6 ab 6 5 384.0 ab 5 5 412.4 a 4 5 427.6 a 3 5 414.6 a 2 5 414.4 a 1 5 326.4 ab Means followed by same letter are not significantly different
Table 13: Plate Organs on Polistes dorsalis by Flagellomere
Flagellomere N Mean 10 5 222.6 b 9 5 250.4 ab 8 5 259.8 ab 7 5 276.2 a 6 5 278.0 a 5 5 286.0 a 4 5 258.2 ab 3 5 247.8 ab 2 5 229.8 b 1 5 172.4 c Means followed by same letter are not significantly different
28
Table 14: Plate Organs on Monobia quadridens by Flagellomere
Flagellomere N Mean 10 5 264.0 c 9 5 381.8 b 8 5 484.6 ab 7 5 519.6 a 6 5 538.0 a 5 5 514.4 a 4 5 539.4 a 3 5 534.2 a 2 5 471.8 ab 1 5 386.4 b Means followed by same letter are not significantly different
Table 15: Total Sensilla Campaniformia by Species
Species N Mean Monobia quadridens 5 11.6 a Polistes metricus 5 16.6 a Polistes dorsalis 5 15.0 a Means followed by same letter are not significantly different
Table 16: Sensilla Campaniformia on Polistes metricus by Surface
Surface N Mean Medial 5 4.2 b Lateral 5 12.4 a Means followed by same letter are not significantly different
Table 17: Sensilla Campaniformia on Polistes dorsalis by Surface
Surface N Mean Medial 5 7.8 a Lateral 5 7.2 a Means followed by same letter are not significantly different
29
Table 18: Sensilla Campaniformia on Monobia quadridens by Surface
Surface N Mean Medial 5 3.6 a Lateral 5 8.0 a Means followed by same letter are not significantly different
Table 19: Sensilla Campaniformia on Polistes metricus by Flagellomere
Flagellomere N Mean 10 5 3.0 a 9 5 1.8 a 8 5 2.2 a 7 5 1.4 a 6 5 1.4 a 5 5 1.4 a 4 5 1.6 a 3 5 0.6 a 2 5 1.4 a 1 5 1.8 a Means followed by same letter are not significantly different
Table 20: Sensilla Campaniformia on Polistes dorsalis by Flagellomere
Flagellomere N Mean 10 5 4.0 a 9 5 2.2 b 8 5 1.6 b 7 5 1.2 b 6 5 2.0 b 5 5 1.6 b 4 5 0.8 b 3 5 0.2 b 2 5 0.8 b 1 5 0.6 b Means followed by same letter are not significantly different
30
Table 21: Sensilla Campaniformia on Monobia quadridens by Flagellomere
Flagellomere N Mean 10 5 1.0 b 9 5 3.8 a 8 5 1.4 b 7 5 2.0 b 6 5 1.6 b 5 5 1.0 b 4 5 0.6 b 3 5 0.0 b 2 5 0.0 b 1 5 0.2 b Means followed by same letter are not significantly different
Table 22: Total Pit Organs by Species
Species N Mean Monobia quadridens 5 174.8 a Polistes metricus 5 92.8 b Polistes dorsalis 5 73.6 b Means followed by same letter are not significantly different
Table 23: Pit Organs on Polistes metricus by Surface
Surface N Mean Medial 5 27.0 b Lateral 5 65.8 a Means followed by same letter are not significantly different
Table 24: Pit Organs on Polistes dorsalis by Surface
Surface N Mean Medial 5 34.4 a Lateral 5 39.2 a Means followed by same letter are not significantly different
31
Table 25: Pit Organs on Monobia quadridens by Surface
Surface N Mean Medial 5 70.0 a Lateral 5 104.8 a Means followed by same letter are not significantly different
Table 26: Pit Organs on Polistes metricus by Flagellomere
Flagellomere N Mean 10 5 7.0 cde 9 5 11.8 abc 8 5 17.4 a 7 5 11.8 abc 6 5 14.2 ab 5 5 10.2 bcd 4 5 8.8 bcde 3 5 4.8 de 2 5 3.6 e 1 5 3.2 e Means followed by same letter are not significantly different
Table 27: Pit Organs on Polistes dorsalis by Flagellomere
Flagellomere N Mean 10 5 5.6 de 9 5 12.6 ab 8 5 14.2 a 7 5 9.2 bcd 6 5 11.4 abc 5 5 8.0 cd 4 5 5.6 de 3 5 3.2 e 2 5 1.8 e 1 5 2.0 e Means followed by same letter are not significantly different
32
Table 28: Pit Organs on Monobia quadridens by Flagellomere
Flagellomere N Mean 10 5 15.8 ab 9 5 32.2 a 8 5 31.2 a 7 5 29.2 a 6 5 19.6 ab 5 5 18.4 ab 4 5 13.4 ab 3 5 8.0 b 2 5 5.4 b 1 5 1.6 b Means followed by same letter are not significantly different
33
CHAPTER VII
FIGURES
Figure 1 Monobia quadridens antenna.
Figure 2 Polistes dorsalis antenna.
Figure 3 Polistes metricus antenna.
34
Figure 4 Significant differences in s. basiconica A by flagellomere. M. quadridens.
Figure 5 Significant differences in s. basiconica A by flagellomere. P. dorsalis.
Figure 6 Significant differences in s. basiconica A by flagellomere. P. metricus. 35
Figure 7 Significant differences in plate organs by flagellomere. M. quadridens.
Figure 8 Significant differences in plate organs by flagellomere. P. dorsalis.
Figure 9 Significant differences in plate organs by flagellomere. P. metricus.
36
Figure 10 Significant differences in pit organs by flagellomere. M. quadridens.
Figure 11 Significant differences in pit organs by flagellomere. P. dorsalis.
Figure 12 Significant differences in pit organs by flagellomere. P. metricus.
37
Figure 13 Significant differences in s. campaniformia by flagellomere. M. quadridens.
Figure 14 Significant differences in s. campaniformia by flagellomere. P. dorsalis.
Figure 15 Significant differences in s. campaniformia by flagellomere. P. metricus.
38
A B
C Figure 16 Plate Organs
NOTE: Plate organ of M. quadridens (A), Plate organ of P. dorsalis (B), Plate organs of P. metricus (C) (Lane et al. 1988).
39
A B
C Figure 17 Sensilla Basiconica A
NOTE: Sensillum basiconicum A of M. quadridens (A), Sensillum basiconicum A of P. dorsalis (B), and Sensillum basiconicum A of P. metricus (C) (Shimizu 2000).
40
A B
C Figure 18 Sensilla Basiconica B
NOTE: Sensilla basiconica B of M. quadridens (A), Sensillum basiconicum B of P. dorsalis (B), Sensillum basiconicum B of P. metricus (C).
41
A B
C Figure 19 Sensilla Campaniformia
NOTE: Sensillum campaniformium of M. quadridens (A), Sensilla campaniformia of P. dorsalis (B), and Sensillum campaniformium of P. metricus (C) (Alm and Kurczewski 1982).
42
A B
C Figure 20 Pit Organs
NOTE: Pit organ of M. quadridens (A), Pit organs of P. dorsalis (B), and Pit organs of P. metricus (C) (Shimizu 2000).
43
A B
C Figure 21 Sensilla Trichodea A
NOTE: Sensillum trichodeum A of M. quadridens (A), Sensillum trichodeum A of P. dorsalis (B), and Sensillum trichodeum A of P. metricus (C) (Lane et al. 1988).
44
A B
C Figure 22 Sensilla Trichodea B
NOTE: Sensillum trichodeum B of M. quadridens (A), Sensillum trichodeum B of P. dorsalis (B), and Sensillum trichodeum B of P. metricus (C) (Lane et al. 1988).
45
A B
C Figure 23 Sensilla Chaetica
NOTE: Sensillum chaeticum of M. quadridens (A), Sensillum chaeticum of P. dorsalis (B), and Sensillum chaeticum of P. metricus (C) (Lane et al. 1988).
46
A B
C Figure 24 Sensillum Spatulatum and Setae of M. quadridens
NOTE: Sensillum spatulatum of M. quadridens (A) (Shimizu 2000), Setae A of M. quadridens (B), and Setae B of M. quadridens (C) (Lane et al. 1998).
47
A B
C D Figure 25 Setae of P. dorsalis
NOTE: Setae A of P. dorsalis (A), Setae B of P. dorsalis (B), Seta C of P. dorsalis (C), and Seta D of P. dorsalis (D) (Lane et al. 1988).
48
A B
C D Figure 26 Setae of P. metricus
NOTE: Setae A of P. metricus (A), Setae B of P. metricus (B), Seta C of P. metricus (C), and Seta D of P. metricus (D) (Lane et al. 1988).
49
A B
C D Figure 27 Sections of Sensilla of P. dorsalis
NOTE: Cross-section of sensillum basiconicum A (A) (Martini et al. 1986), Cross-section of plate organ (B), Longitudinal section of plate organ (C) (Bleeker et al. 2004), and Section of pit organ (D) (Lacher 1964).
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