Males usually had the body size smaller than that of females. The male much often
flew around the nesting site, when it found female, immidiately grasped the female from the
back. It used its mandible to bite the pronotum, placing his forelegs on both sides of her
mesosoma just anterior to her hindlegs, and his midlegs and hindlegs straddling her abdomen.
The male tapped his antennae on the top of her head while probing the apex of her abdomen
with his genital claspers, then bent his abdomen and attempted to clasp the female’s genitalia
from above. The male mated the female continously. During the copulation they often buzzed
loudly. We did not know whether or not the buzz that was from the male or the female made.
The copulation time varied between 20 second to 8 minutes. After completing the couple,
immidiately both the male and female flew away
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uals number of specimens collected by the number total of all
specimens collected:
Number of specimens collected
Rate (%) = x 100
The number total of all specimens collected
8
2.3.1.2. Methods of pinning specimens
Specimens were pinned and dried and then kept in insect boxes and deposited at the
Institute of Ecology and Biological Resources, Vietnam Academy of Science and technology,
18 Hoang Quoc Viet, Cau Giay, Ha Noi.
2.3.1.3. Methods of dertermination
We used microscopes Olympus model SZ60 to observe morphological characters of
specimens and publications with keys to determine the scientific name of species, such as:
Kirby (1798), Bingham (1897), William (1919), Menke (1961) [100], Tsuneki (1963), Menke
and Willink (1964), van der Vecht and van Breugel (1968), Bohart and Menke (1976),
Krombein (1984), Hensen (1987, 1988, 1991), Ohl and Höhn (2011), Danilov (2012, 2015,
2018), Dollfuss (2008, 2013, 2016), Kim et al. (2014), Kumar et al. (2014) and Dörfel and
Ohl (2015). Terminology used in the thesis followed that of Bohart and Menke (1976).
Determination of subspecies of S. madraspatanum was based on publications of van
der Vecht and van Breugel (1968) and Bohart and Menke (1976). The key was based on that
of van der Vecht and van Breugel (1968).
2.3.1.4. Methods of study on the distribution of sphecid wasps of the family Sphecidae
+ At areas of the study, 4 habitats choosen were natural forest habitat, artificial forest
and orchard habitats, shrub habitat, and resident habitat.
2.3.2. Methods of study on biological characteristics of S. madraspatanum
Under the laboraroty condition, the average temperature 28,1 – 28,6C and the average
humidity 80,9 – 81,2%. Research methods on biological characteristics of S. madraspatanum
were based on those of Krombein (1984), Polidori et al. (2005) and Gess và Gess (2014).
2.3.2.1. Methods of study on the nesting construction
New nests of S. madraspatanum collected in the field were splited to take nesting
contents including: Nesting materials, nesting colours, shape of cells and nests, number of
cells and number of nesting floors. Cell and nest sizes were measured by mini meter rulers.
Nests of S. madrapatanum were distinguished from those of species of the genus
Sceliphron by having adults that were emerged or pupae or and adults that had been died into
the nest. S. madrapatanum and its subspecies were determined based on the key of van der
Vecht and van Breugel (1968).
2.3.2.2. research methods of the developmental time of stages
Stages of S. madraspatanum consisted of: egg, larva, prepupa, pupa and adult. We used
microscope model SZ60 to observe and to describe morphological characterictics of these
stages. The size of egg stage was measured by a magnifier attached with a mini meter ruler,
and that of the others was measured by a mini meter ruler. Prepupae and pupae were romoved
from their cocoons. The length was measured from the head to the end part of abdoment. The
width of the egg and larva stages was taken at the center of the body, that of the prepupa stage
was hold at the widest ternite, and that of the pupa and adult stage was measured between
tegulars on the mesoscutum. Pictures were taken using Canon model SD3500 IS.
Newly built nests of S. madraspatanum in the field were spilted to take nesting contents
such as: eggs, larvaecocoons, adults, and prey. Eggs were collected from nesting cells
provisioned in the same day. The contents were then put in small glass tubes (5 - 7 cm long
and 0,6 - 1 cm in diameter). Because the immature stages of the wasp in the field is into mud
nests without light, the contents were also put in the same condition (into carton boxes). Each
of them was attached to a small ticket (5 cm long and 3 cm wide) that aimed to write down
information of results. These contents were observed every day. Tubes contained eggs and
larvae were looked at with the microscope of the magnification of 40 times. In each of these
stages, 20 – 25 individuals were kept into Ethanol 95% for study on morphology..
9
After emerging, adults of S. madraspatanum were reared into insect net cages measured
by 50 x 50 x 50 cm and food of bee’s honey diluted to 50% were provided to them. Wet mud
and prey were also put into theses cages. We wrote down notes related to completion time of
a nesting cell and methods of provision and them dissected these cells to find eggs newly laid.
These eggs would be foundation to determine time before laying of females of the wasp.
Another method to determine this time was that we dissected adult females after of 4 – 7 days
of age to find their eggs. The eggs with more darker colour than were looked at to know
whether or not they could hatch. If the eggs laid, we would determine time before laying of
the females (referrced from Ohl và Linde, 2003).
Temperature and humidity in the laboratory condition were taken using Max-Min
Thermo Hygro, CT-138B.
2.3.2.3. Methods of study on life time of adults
Adults emerged in the laboratory were reared into plastic jars (20 cm high and 12 cm
in diameter) with covers made of mosquito nets. Each jar kept 2 adults either males or females.
The adults were fed with 50%-diluted bee’s honey. A piece of cotton was wetted by the honey
and hang under covers.
The sex was distinghuished by: normal observations and observations under
magnifiers.
2.3.2.4. Mothods of study on life time of immature stages
Methods used here were the same mentioned above (item 2.3.2.2).
Formulas of life rates of immature stages were:
The number total of eggs laid
Life rate of egg (%) = x 100
The number total of eggs tested
The number total of larvae emerged
Life rate of larva (%) = x 100
The number total of larvae tested
The number total of pupae emerged
Life rate of pupa (%) = x 100
The number total of pupae tested
For old nests of S. madraspatanum, each emerged hole determined as successful
emergence of an adult because the holes were made by adults getting arise.
2.3.2.5. Methods of study on sex rate
Newly built nests of the wasp collected in the field were choosen and numberred (from
1 to the end), then put into plastic jars (the same size mentioned above). These jars were
coverred by mosquito nets and were observed every day. When adults emerged, we noted
number marked on the surface of their nest and determined their sexes.
The sex rate in the field was determined by catching adults and then find their sexes.
We used insect nets to catch these adults.
In the laboratory, we determined the sexes of adults immediately after they emerging.
2.3.3. Mothods of study on ecological characteristics of S. madraspatanum
2.3.3.1. Mothods of study on nesing sites
New and old nests of S. madraspatanum were collected at all areas of study in the field.
These nests were numberred, noted on time of collection, location, substrate of nests attached
and highlight of nesting sites. Pictures of nesting sites were taken. In the laboratory, we
analysed and assembled nesting sites of the species.
10
2.3.3.2. Methods of study on overwintering time
New nests of S. madraspatanum collected from October to November were splited to
take cocoons newly completed. The cocoons were put into glass tubes (50 - 80 mm long, 6 -
10 mm in diameter) and these tubes then put in plastic boxes coverred. The boxes were put
under the eaves (the same natural condition). We checked the tubes once a week, and when
we found pupae we determined that time was end of overwintering time. We also surveyed
nests of the wasp in the field at the beginning of summer to remark the beginning time of
occurrence of the species.
2.3.3.3. Methods of study on yearly occurred time of adults
We investigated to find new nests of S. madraspatanum from October to November
between and remarked these nests as well as ending time of action of adults in year. Sites
serveyed were under bridges and desolated houses. The beginning of following April, we
observed these nests, if adults emerged or emerging holes appeared we calculated the time of
occurrence of the wasp in the field.
Cocoons methoded as mentioned in the item of the overwintering time previously.
When adults emerged we calculated the time of occurrence of the wasp in year.
2.3.3.4. Methods of study on the effect of food to life time of adults
We did this study within 4 formulas: no food, fresh water, 50% diluted bee’s honey,
and fruit suger cane. After immediately emerging, adults were reared into plastic jars (20 cm
high and 12 cm in diameter), each with 2 adults. The jars were coverred by morquitor nets.
We checked the jars every day until adults dead.
2.3.3.5. Mmethods of study on prey
We chose 10 nests of S. madraspatanum collected in several different locations of the
areas of study and splited them to take their contents, mainly prey. Prey taken were put into
jars contained ethanol 70%. Prey were then sent to specialists to determine them to genera or
species level.
Occuring rate of prey into nests was showed such as:
The number total of nests contained prey
Occuring rate (%) = x 100
The number total of nests tested
2.3.3.6. Methods of study on predators
Predators of S. madraspatanum were defined by: Observations in the field and analysis
of nests of the wasp in the laboratory.
2.3.3.7. Methods of study on the effect of human activities to nesting sites of S.
madraspatanum
We collected information about the effect of human activities to nesting sites of S.
madraspatanum through interviews, specially checked distubances of human activities to
nests built into houses. Information consisted of: driving away, destroying nest, destroying
nesting site, and cultivating activity.
2.3.4. Methods of study on habits of S. madraspatanum
2.3.4.1. Methods of study on nesting, egg-laying, and mating habits
We chose nesting sites, which many adults of the species appeared, such as: under
bridges and into desolated houses, because it would be easy for us to observe its habits. Sites
chosen here for observations on nesting habits were into a desolated house built in a fruit
garden at Thanh Tan commune, Luong Son district, Hoa Binh province; a desolated house at
Lien Mac commune, Bac Tu Liem district, Ha Noi city; into a desolated house at Hong Minh
commune, Hung Ha district, Thai Binh province. Sites chosen for observations on mating
11
habits were into a desolated house at Hong Minh commune, Hung Ha district, Thai Binh
province and under bridge at Co Nhue precinct, Bac Tu Liem district, Ha Noi city.
When females of the wasp began to build its nest, we observed them, wrote down notes
on methods of making pieces of mud, bringing mud, and building nest with mud... and took
pictures using Canon model SD 3500IS to aim analysis more than data. Nest-building time
was taken by a Stopwatch model SPT-20. With cells that have not yet completed in the day,
we used a violet inked pen to remark onto those cells and took a continuingobservations in
the following day.
Methods of study on egg-laying habits of the wasp were based on analysis of nests in
which cells contained eggs and prey into.
2.3.4.2. methods of study on feeding habits of larva
Feeding and cocooning habits of larvae were observed based on methods of study on
the developmental time of immature stages (also see the item 2.3.2.2).
2.3.4.3. Methods of study on emerging habits of adult
In the laboratory: Fresh nests of S. madraspatanum collected in the field were put
plastic jars (12 cm high, 10 cm in diameter) with covers of morquito nets. The nests were
observed every day, if when high-intensitied buzzes were made by adults, we observed and
wrote down notes on emerging habits of the species.
In the field: Fresh nests of S. madraspatanum built under a bridge at Co Nhue precinct,
Bac Tu Liem district, Ha Noi city were observed. To take the emerging time of adults, we had
based on the nest-built time and marking nest cells.
2.3.5. Methods of study on analysing data
Data yielded in the project were analysed using the Data Analysis in Microsoft Office
Excel 2007, and ANOVA.
CHAPTER 3.
RESULTS AND DISCUSSION
3.1. The composition of solitary wasps of the family Sphecidae and their distribution in
several provinces of the northwest part of Vietnam
3.1.1. The composition of solitary wasps of the family Sphecidae
The composition of solitary wasps of the family in several provinces of the northwest
part of Vietnam was studied and showed in table 3.1. A total of 25 species and 6 subspecies
belonging to 7 genera into 4 subfamilies were recorded. Of which, C. tanvinhensis Pham and
Ohl, 2019 had decribed already, 4 species and 3 subspecies were recorded for the sphecid
wasp fauna of Vietnam, and 9 species and 6 subspecies were recorded for the first time from
in northwest Vietnam.
Table 3.1. The composition of solitary wasps of the family Sphecidae in several
provinces of the northwest part of Vietnam
No. Species name Rate of
collection (% )
A Subfamily Ammophilinae
I Genus Ammophila
1 Ammophila clavus (Fabricius, 1775) 15.87
2 Ammophila globifrontalis Li và Yang, 1995** 0.55
3 Ammophila laevigata Smith,1856 5.47
B Subfamily Chloriontinae
II Genus Chlorion
4 Chlorion lobatum (Fabricius, 1775) 2.55
12
C Subfamily Sceliphrinae
III Genus Chalybion
5 Chalybion bengalense (Dahlbom, 1845) 12.59
6 Chalybion dolichothorax (Kohl, 1918) 0.18
7 Chalybion gracile Hensen, 1988*** 0.90
8 Chalybion japonicum (Gribodo, 1883)*** 9.30
9 Chalybion malignum (Kohl, 1906)** 6.02
10 Chalybion tanvinhensis Pham và Ohl, 2019* 0.54
IV Genus Sceliphron
11 Sceliphron deforme (Smith, 1856) 4.56
12.1 Sceliphron javanum petiolare Kohl, 1918*** 1.28
12.2 Sceliphron javanum chinensis van Breugel, 1968*** 0.36
13 Sceliphron madraspatanum Fabricius, 1781 20.43
13.1 Sceliphron madraspatanum sutteri van der Vecht, 1957** 0.54
13.2 Sceliphron madraspatanum andamanicum Kohl, 1918** 0.54
13.3 Sceliphron madraspatanum conspicillatum (Costa, 1864)** 3.28
13.4 Sceliphron madraspatanum kohli Sickmann, 1894*** 1.82
D Subfamily Sphecinae
V Genus Isodontia
14 Isodontia aurifrons (Smith, 1859)*** 0,90
15 Isodontia chrysorrhoea (Kohl, 1890)*** 0,18
16 Isodontia elsei Hensen, 1991** 0,36
17 Isodontia nigella (Smith, 1856)** 2,37
18 Isodontia sp.1 0,18
19 Isodontia sp.2 0,90
20 Isodontia sp.3 1,82
VI Genus Prionyx
21 Prionyx viduatus (Christ, 1791)*** 1,64
VII Genus Sphex
22 Sphex argentutus Fabricius, 1787 1,46
23 Sphex diabonicus Smith, 1858 0,36
24 Sphex sericeus (Fabricius, 1804) 0,54
25 Sphex subtruncatus Dahlbom, 1843 2,19
Total 100
Remark: *new species, **new record for Vietnam, ***New record in northwest Vietnam
3.1.2. Morphological characteristics of Chalybion tanvinhensis Pham and Ohl, 2019
Specimens examined: 1♀, Tan Vinh, Luong Son, Hoa Binh, 2.v.2017, Phong Huy
Pham; 2♂, Ban Chieu, Muong Thai, Phu Yen, Son La, 11.viii.2017, Phong Huy Pham.
Morphology:
Female (Figs. 1‒6): Body length 20 mm, forewing length 15 mm.
Head (Fig. 1): In frontal view subcircular, about 1.23 times as wide as high. Mandible
with inner subapical tooth. Clypeus moderately convex, without median carina; Anterior
margin of clypeus with five distinct teeth: lateral tooth, separated distantly from submedian
tooth, narrow and sharp; submedian tooth blunt, broader than median and lateral teeth, and as
long as median tooth. Frontal line present. Ocelli not reduced; distance between hindocelli
about equal to that from hindocellus to inner eye margin and about 0.43 times that from
hindocellus to vertex margin. Clypeus coarsely, sparsely punctate. Frons coarsely, irregularly
13
transversely punctatorugose. Vertex and gena finely, sparsely punctate. White setae sparse on
clypeus, denser than on gena. Lower inner orbit area with moderately dense pubescence.
Flagellar ratio (length of the first flagellomere divided by that of the second flagellomere)
0.82; orbital ratio (shortest interocular width across vertex divided by shortest interocular
width across clypeus) 0.83; clypeal ratio (length of clypeus divided by shortest interocular
width across clypeus) 0.82; hypostomal cavity ratio (length of hypostomal cavity divided by
width) 0.89.
Mesosoma (Figs. 2‒5): In dorsal view, width between tegulae equal to that of
pronotum at midlength. Pronotum with notch at middle and series of transverse striae
anteriorly. mesoscutum moderately transversely striate, interspaces shallowly punctate;
mesopleuron coarsely, densely punctate. Scutellum, metanotum, and upper metapleural area
rather coarsely, densely punctate. Lower metapleuron conspicuously, transversely striate.
Dorsal propodeum coarsely, transversely striate; side of propodeum punctate-reticulate.
Mesosoma ratio (length of mesosoma divided by height) 2.18.
Metasoma (Figs. 5‒6): Integument smooth, shiny, with silvery pubescence. Petiole
conspicuously curved. Metasomal sternum IV without micropubescence. Petiole much
shorter than hindbasitarsus. Petiole-basitarsal ratio (length of petiole divided by that of
hindbasitarsus) 0.70.
Color: Integument dark blue. Mid and hind legs, propodeum and metasoma with
violaceous reflections; antenna black except scape and pedicel dark blue. Mandible, labial
palpus and maxillary palpus black. Forewing slightly, rather unevenly brownish, medial and
submedial cells mostly uniformly hyaline, infuscate at apex (Figs. 3‒4). Hindwing hyaline,
somewhat infuscate at apex (Fig. 4).
Hình 3.1 - 3.6: Chalybion tanvinhensis Pham và Ohl, 2019 (female)
3.1. Head, frontal view. 3.2. Mesosoma, dorsal view. 3.3. Right forewing. 3.4. Habitus,
dorsal view. 3.5. Habitus, lateral view. 3.6. Metasoma, sternum view.
Male (Figs. 7‒10): Body length 14.5-15.5 mm, forewing length 10.5 mm.
Structure as in female, but differing as follows:
Head (Fig. 7): Mandible without inner subapical tooth. Clypeus without teeth, median
and submedian tooth fused into single protruding lobe. Distance between hindocelli 0.62-0.67
times that from hindocellus to inner eye margin and about 0.31-0.33 times that from
hindocellus to vertex margin; antenna with placoids on seventh to ninth flagellomeres.
Flagellar ratio (length of the first flagellomere divided by that of the second flagellomere)
0.89-0.94; orbital ratio (shortest interocular width across vertex divided by shortest
interocular width across clypeus) 0.93-0.97; clypeal ratio (length of clypeus divided by
14
shortest interocular width across clypeus) 0.60-0.67; hypostomal cavity ratio (length of
hypostomal cavity divided by width) 0.89-0.96.
Mesosoma (Figs 8‒9): Mesopleuron coarsely, densely punctatorugose. Posterior
propodeal surface coarsely, densely punctate-reticulate. Mesosoma ratio (length of mesosoma
divided by height) 2.18-2.21.
Metasoma (Figs. 9‒10): Metasomal sterna IV and V with large patch of
micropubescence. Petiole-basitarsal ratio (length of petiole divided by that of hindbasitarsus)
0.58-0.64.
Hình 3.7 - 3.10: Chalybion tanvinhensis Pham và Ohl, 2019 (male)
3.7. Head, frontal view. 3.8. Habitus, dorsal view. 3.9. Habitus, lateral view. 3.10.
Metasoma, sternum view.
Distribution: Vietnam: Hoa Binh, Son La
Chalybion tanvinhensis belonged to the C. fabricator species group. This group has 8
species having been described hitherto: C. malignum (Kohl), C. accline (Kohl), C. magnum
Hensen, C. sulawesii Ohl, C. lividum Hensen, C. fabricator (Smith) và C. hainanense
Terayama and Tano, and C. tanvinhensis Pham and Ohl.
3.1.3. Distribution of solitary wasps of the family Sphecidae at the areas of study
Distribution of solitary wasps of the family Sphecidae at 4 habitats was presented in
table 3.3. Said results showed 19 species belonging to 7 genera in 4 subfamilies being
recorded in the natural forest habitat, 25 species belonging to 7 genera in 4 subfamilies in the
artificial forest and orchard habitats, 17 species belonging to 4 genera in 4 subfamilies in the
shrub habitat, and 10 species belongting to 4 genera in 4 subfamilies in the resident habitat.
Table 3.3. Distribution of solitary wasps of the family Sphecidae in habitats of study
No.
Species name
Habitat
Natural
forest
Artificial
forest
and
orchard
Shrub Resident
1 Ammophila clavus + + + +
2 Ammophila globifrontalis - + - -
3 Ammophila laevigata + + + +
4 Chlorion lobatum + + + +
5 Chalybion bengalense + + + +
6 Chalybion dolichothorax + - - -
7 Chalybion gracile - + - -
8 Chalybion japonicum + + + +
9 Chalybion malignum + + + -
10 Chalybion tanvinhensis - + - +
11 Sceliphron deforme + + + -
12.1 Sceliphron javanum petiolare + + - -
12.2 Sceliphron javanum chinensis + + - -
15
13 Sceliphron madraspatanum + + + +
13.1 Sceliphron madraspatanum sutteri - - + +
13.2 Sceliphron madraspatanum
andamanicum
- + + -
13.3 Sceliphron madraspatanum
conspicillatum
- - + -
13.4 Sceliphron madraspatanum kohli - + + +
14 Isodontia aurifrons + + - -
15 Isodontia chrysorrhoea + - - -
16 Isodontia elsei - + - -
17 Isodontia nigella + + + -
18 Isodontia sp.1 - - + -
19 Isodontia sp.2 + + - -
20 Isodontia sp.3 + + - -
21 Prionyx viduatus + + + -
22 Sphex argentutus + + + +
23 Sphex diabonicus + + - -
24 Sphex sericeus - + + -
25 Sphex subtruncatus + + + -
Tổng 19 25 17 10
Remark: + present, - absent
3.2. Some biological characteristics of S. madraspatanum
3.2.1. nesting construction
Materials of nesting construction of S. madraspatanum were mud or coal. The coulour
of the nest was based on that of mud, but that of nests known mainly was brown.
The nest of the wasp was constructed from 1 to 17 cells. Whereas nests constructed
only one cell, no cover was made, but nests with more than 2 cells, a cover was made. Based
on nesting sites and nesting substrates, nesting covers thickened variously from
2.5 to 10.5 mm. The outside surface of nests was often rough, but that of some was flat
and some nests were with several mud thorns on the nesting surface.
On many instances, there were space between pieces of mud put on the nesting
surface. Nesing cells were pipe-shaped and round ends. Partitions between nesting cells
measured 1.0 – 1.2 mm, average 1.1 0.07 mm. The outside surface of cells was rough,
but the into surface was smooth. Nesting cells built adjoining, and partition of one cell
became that of the next cell. There was a space between cells, but this space was made fully
with mud. Nests of the wasp were usually constructed from 1 – 3 floors, number of cells per
floor decreased following floors above. Cell size contained female was often larger than
that contained male (table 3.4).
Table 3.4. Nesting cell size of S. madraspatanum
Length (cm) (n = 32) Width (cm) (n = 32)
Female Male Female Male
2.7 – 3.1
2.86 0.12a
2.4 – 2.8
2.63 0.10b
0.7 – 0.9
0.82 0.06c
0.5 – 0.75
0.66 0.07d
Remard: In limit of values brough different letters showing confident variousness at
probability P 0.05, n was nesting cells tested
16
3.2.2. The developmental time of stages
+ Egg stage: Egg of S. madraspatanum was white of yellowish, with egg cover smooth
and shiny. Eggs were saussage- shaped, round at the two ends, and moderately curved. The
developmental time of egg varied 2 - 3 days, average 2.23 0.43 days (table 3.6).
+ Larva stage: The larvae of S. madraspatanum had three instars. In general, the
coulor of larva was yellowish. Dorsal legs were short and coloured yellow. There were clear
joints on their body, each bringing one breathing orifice. Skin of larvae smooth. The back
of larva was with a ridge longed from the first joint to the anus joint. The developmental
time of the 1st instar larva was 1-2 days, average 1.39 0.49 days; the 2st instar larva from
1 - 2 days, average 1.65 0.48 days; the 3st instar larva from 4 - 5 days, average 4.31
0.47 days (table 3.6).
+ Pupal stage: The pupa of S. madraspatanum was free and was into a cocoon coloured
brownish. There were 2 stages: the prepupa and the pupa. The first was cloured yellow and
the body had 13 joints ranged from the thorax to abdomen (3 thorax joints and 10 abdomen
joints), the largest 6 – 8 joints. The last had morphological body similar to that of adult,
excepting clour of pupae newly emerged and thorns distributed the two sides of the body. The
developmental time of the prepupa was 5 - 6 days, average 5.58 0.50 days. The
developmental time of the pupa was 13 - 16 days, average 14.48 1.14 days (
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