A New Eustenogater Species (Hymenoptera: Vespidae; Stenogastrinae), the First Hover Wasp Known to Overwinter on the Nest

2006·0.43 MB·English

by  Fuki Saito

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3534, 11 pp., 15 figures September 08, 2006
A New Eustenogater Species (Hymenoptera:
Vespidae; Stenogastrinae), the First Hover Wasp
Known to Overwinter on the Nest
FUKI SAITO,1 LIEN P. T. NGUYEN,2 JAMES M. CARPENTER,3 AND
JUN-ICHI KOJIMA1,4
ABSTRACT
A new species of hover wasp, Eustenogaster nigra Saito and Nguyen, is described based on
females and males collected mainly in northern Viet Nam. The wasp is also distributed in
mountainousareas of central Viet Nam andthe southern part of China. Alldistribution records
arefromareaswithmore-or-lessdistinctseasonsintermsoftemperature.Thenestisdescribedand
compared with those of other Eustenogaster species. A nest collected during early spring in Tam
DaoNationalParkinnorthernVietNam,wherethereisamore-or-lessdistinct‘‘winter’’,hadfive
malesand sixvirgin females, suggestingthatboth sexes overwinteron theirnest.
INTRODUCTION Stenogastrinae, however, has been made since
Not only because the hover wasps, CarpenterandKojima(1997)pointedoutthat
Stenogastrinae, exhibit considerable diversity the stenogastrine taxonomyhad notbeen well
insociallifeandnestingbehavior(reviewedin studiedcomparedtotheothertwosocialwasp
Turillazzi, 1991), but also the group is the subfamilies, Vespinae and Polistinae.
basal clade of the social wasps (Carpenter, Sevengeneraarecurrentlyrecognizedinthe
1991), the subfamily is the key group for subfamily (Carpenter, 2001), and the late
understanding of social evolution in the J. van der Vecht had been preparing taxo-
wasps. Little progress in the taxonomy of the nomic revisions of all the genera of the
1Natural History Laboratory, Faculty of Science, Ibaraki University, Mito, 310-8512 Japan (F.S., nd5507t@mcs.
ibaraki.ac.jp;J.K.,[email protected]).
2Insect Ecology Department, Institute of Ecology and Biological Resources, Vietnamese Academy of Science and
Technology,18HoangQuocVietRoad,NghiaDo,CauGiay,HaNoi,VietNam([email protected]).
3DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory,([email protected]).
4DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory.
CopyrightEAmericanMuseumofNaturalHistory2006 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3534
Stenogastrinae; however, he died in 1992 describe the nests and some biological aspects
before having completed the revisions of of this new Eustenogaster species in Tam Dao
Eustenogaster van der Vecht, 1969, National Park, in the northern part of Viet
Liostenogaster van der Vecht, 1969, and Nam, where social wasps in general appear to
Parischnogaster von Schulthess, 1914. The have a winter resting period.
revisions of these genera that van der Vecht
began were taken over by C.K. Starr on MATERIALS AND METHODS
Parischnogaster and by S. Turillazzi on
Liostenogaster, but descriptions of only Most of the Vietnamese specimens were
Liostenogaster species have appeared collectedbyourselvesandaredepositedinthe
(Turillazzi, 1988, 1999; Turillazzi and Carfi, InstituteofEcologyandBiologicalResources,
1996). Hanoi, Viet Nam (IEBR), and as long-term
The late van der Vecht distributed speci- loans in the American Museum of Natural
mens of social wasps labeled with his manu- History (AMNH) and the Natural History
script names to various collections, and this Collection, Ibaraki University, Mito, Japan
was also the case for the Stenogastrinae. (IUNH). Some specimens that we did not
A regrettable consequence of this is that collect are also in the IEBR.
some of these names have been published Morphologicalaswellascolorcharactersof
adult wasps were observed on pinned-and-
as nomina nuda, including one name of
dried specimens under a stereoscopic dissect-
Ropalidia Gue´rin-Me´neville, 1831 (subfamily
ing microscope. Male genitalia were dissected
Polistinae) and two Liostenogaster names
out, briefly cleared in 10% KOH, and ob-
(Carpenter, 1982; Hansell, 1987a). The
served in glycerin under a stereoscopic dis-
Ropalidia name has been validated (Kojima,
secting microscope. Nest measurements were
1989) and the two Liostenogaster species were
made with digital calipers for envelope and
subsequently described under different names
cell sizes and with a micrometer for the cell
(Turillazzi,1988,1999;seealsoKojima,2006).
wall thickness.
Recently,oneofvanderVecht’smanuscript
Abbreviations other than museum/institu-
names in Eustenogaster was published as
tional acronyms are as follows: F.S., Fuki
a nomen nudum. Nguyen and Khuat (2004)
Saito; ISD-c, collectors of the Insect
published the name in quotation marks, with
Systematic Department of the IEBR; J.K.,
a note that it came from van der Vecht’s
Jun-ichiKojima;J.M.C.,JamesM.Carpenter;
unpublished notes, and they gave its diagnos-
NP, National Park; N.L., Lien Thi Phuong
tic characters in a key, together with a figure
Nguyen.
of the head in frontal view and a list of
specimens examined. There was no intention
DESCRIPTION OF NEW SPECIES
to make the name available, as it is not under
the current International Code of Zoological
Eustenogaster nigra Saito and Nguyen,
Nomenclature (ICZN, 1999). That is, Nguyen
new species
and Khuat (2004) neither made an explicit
statementthatthenameisnew(article16)nor
fixed the name-bearing type (article 16.4). In Eustenogasterscitula(Bingham,1897):Nguyenand
the present paper, we validate the name. Khuat, 2003: 696[misidentification, partly].
Eustenogaster is distributed from India to Eustenogaster nigra: Nguyen and Khuat, 2004: 38,
Southeast Asia, mainly in tropical areas. The 40. Nomen nudum.
nesting biology has been described for several FEMALE: Bodylength(head+mesosoma+
species (Williams, 1919; von Schulthess, 1927; first two terga) 19.5–21.5 mm (holotype:
Bell, 1936; Pagden, 1958; Sakagami and 19.5 mm); forewing length 14.0–15.2 mm (ho-
Yoshikawa, 1968; Yoshikawa et al., 1969; lotype:15.2 mm).Headinfrontalview(fig. 1)
Krombein, 1976; Ohgushi and Yamane, 1983; broad, about 1.1 times wider than high. Eyes
Ohgushi et al., 1983, 1986, 1988, 1990; enlarged, in lateral view strongly swollen at
Hansell, 1987b; Turillazzi, 1991; Francescato least in ventral half (fig. 2), maximum width
et al., 2002). In the present paper, we also about 4.5 times as wide as that of gena.
2006 SAITO ETAL.: NEW HOVERWASP 3
Clypeus convex, with apex sharply pointed Genitalia: Parameral spines not dilated,
triangular;supraclypeal areairregularly punc- with distinct depression at the base (fig. 9).
tuate, with small impunctate, flat area below Volsella bent at anterior margin, with digitus
anterior ocellus. Anterior ocellus slightly beak-shaped (fig. 10). Aedeagus slender, in
elliptical in shape (0.32–0.36 mm wide, 0.28– lateral view slightly dilated apically (fig. 11),
0.30 mm long), about 1.4 times larger in withapairofprojectionslocatedlaterobasally
diameter than more-or-less circular posterior (fig. 12).
ocelli (diameter 0.22–0.26 mm); distance be- U
tween inner eye margin and posteriorocellias TYPEMATERIAL:Holotype labeled‘‘VietNam:
Tam Dao (outside town), ca. 900m, Vinh Phuc,
longaswidthofanteriorocellus;ocellicloseto
7.iii.2005, L.T.P. Nguyen, F. Saito & J. Kojima,
each other (fig. 3); distance between anterior
Nest #VNM-S2005B’’ and ‘‘No. 8’’, in IEBR; on
and posterior ocelli shorter than diameter of long-term loanto IUNH.
posterior ocellus; posterior ocelli separated Paratypes (unless the depository is mentioned,
from each other by about distance equal to theparatypesaretentativelyinIUNHonlong-term
-
their diameter. Mandible with three teeth; loanfromIEBR):CHINA:2 (AMNH),Yen-ping
- -
proximal tooth small,sharply pointed; middle [5 Nanping or Yanping], [1 , 31.viii.1917; 1 ,
U
and distal teeth extended, bluntly projected. 9.vi.1917].VIETNAM:ThaiNguyenProvince:1 ,
-
Cat Ne, Dai Tu, 23.x.2004, ISD-c; 11 , Than Xa,
Scutumfinely,denselypunctate(interspaces - -
Vo Nhai, ISD-c [7 , 470m, 16.x.2004; 2 , 90m,
smaller than punctures), with median carina -
19.x.2004;2 ,70m,17.x.2004];PhuThoProvince:
developed anteriorly.Scutellumhairy,densely - -
32 , Xuan Son NP, N.L. [6 , 400m, 11–
punctate, strongly convex medially, separated - -
16.vi.2004; 8 , 500m, 12.vi.2004; 18 , 600m,
from metanotum by deep but narrow furrow. 13.vi.2004]; Vinh Phuc Province: 5U, 21-, Tam
- -
Metanotum rather strongly convex, hairy, Dao NP [1 , 800m, 7.ix.2000, N.L.; 1 , 1000m,
- -
rugosely punctate in posterior margin. 08.ix.2000,N.L.;1 ,800m,12.v.2003,N.L.;13 ,
U -
Mesepisternum densely punctate at least in 800m,01–04.vii.2003,N.L.;5 ,5 ,samedataas
-
dorsal half. Propodeum hairy, with shallow, holotype; 5 (IEBR), 500–700m, 20.viii.2005,
-
J.K.]; 1 , Ngoc Thanh, Me Linh, 100m,
small punctures. U -
24.v.2000, N.L.; 1 , 1 , Ngoc Thanh, Me Linh,
First metasomal tergum 8.0–8.6 mm long, -
22.viii.2000, X.L. Truong; 3 , Tay Thien Mt.,
about 6.1 times as long as maximum width, U -
26.viii.2004, N.L.; Ha Tay Province: 1 , 13 , Ba
and 7.2 times longer than height; second - U -
ViNP,N.L.[12 ,1 ,800m,18.ix.2000;1 ,400–
tergum strongly convex dorsally (fig. 4); sixth 600m, 02.vi.2001]; 3-, Yen Bai, Ba Vi, .100m,
tergum with a small tubercle (fig. 7). 01.vi.2001, N.L.; Hai Phong Province: 5-, Cat Ba
-
Color: Body black; head with no spots or NP, 15–18.vii.2003, N.L.; Hoa Binh Province: 1 ,
markings; paired short lines along poster- PaCo,MaiChau,1100m,23.iv.2002,V.T.Hoang;
-
odorsalmarginofpronotum,andscrobalspot Nghe An Province: 3 , Chau Cuong, Quy Hop,
- - U -
X.H.Le[2 ,17.vii.2004;1 ,14.vii.2004];1 ,1 ,
and spot below it (both often reduced) on
Mon Son, Con Cuong, 22–24.vii.2004, N.L.; Ha
mesepisternum yellow; posterior two-thirds of - U
TinhProvince:1 ,1 (AMNH),17kmSE,Huong
firsttergumdarkreddishbrown;thirdtergum
Son, 18u229N, 105u139E, 180m, 19–23.iv.1998,
anterolaterallywithpaired,short,yellowlines. -
JMC; 1 (AMNH), Huong Son, 200–300m,
Legsdarkbrowntoblack;femorapalebrown 18u219N, 106u159E, 4.v.1998, J.M.C., Long,
apically. Wings semihyaline, pale brown, Grimaldi, Herman, and Silva; 1U, 4-, Son Hong,
darker along anterior margin of forewing; HuongSon,25.iv.2004,X.L.Truong.
veins brown. OTHER SPECIMENS EXAMINED: VIETNAM:Lang
-
MALE: Body length (head + mesosoma + Son Province: 2 , Bac Son,-01.vii.2003, X.L.
first two terga) 18–20 mm; forewing length Truong; Vinh Phuc Province: 2 , Tam Dao NP,
-
14.5–15.5 mm. Structure and coloration sim- 800m,01–04.vii.2003,N.L.;HoaBinhProvince:1 ,
-
LacThinh,YenThuy,30.iv.2002,D.L.Khuat;1 ,
ilartofemale,butclypeuslessconvexandwith
PaCo,MaiChau,1100m,24.iv.2002,D.L.Khuat;
apexbluntlyangled,sometimeswithamedian - -
3 ,DaPhuc,YenThuy,3.v.2002,V.T.Hoang;1 ,
yellowmarkofvariablesize;mandibularteeth
Pa Co, Mai Chau, .1100m, 21–23.x.2002, D.L.
rounded apically (fig. 8); second tergum less Khuat; Ninh Binh Province: 3-, Cuc Phuong NP,
stronglyconvexdorsally(fig. 5);tergawithno NhoQuan,7.v.2002,V.T.Hoang;9-,CucPhuong
tubercle. NP,NhoQuan,08.v.2002,V.T.Hoang.
4 AMERICAN MUSEUMNOVITATES NO. 3534
Figs.1–12. Eustenogasternigraholotype andparatypes.Scalebarsare1 mm.1–3. Eustenogasternigra
Saito and Nguyen, n.sp., female holotype. Head. 1. Frontal view. 2. Lateral view. 3. Dorsal view. 4–6.
Secondmetasomalterguminlateralview.4.EustenogasternigraSaitoandNguyen,n.sp.,female.5.E.nigra,
male.6.E.scitula,female.7.EustenogasternigraSaitoandNguyen,n.sp.,femaleparatype.Sixthmetasomal
2006 SAITO ETAL.: NEW HOVERWASP 5
ETYMOLOGY: The specific name is the wide)oflighteranddarkerbrownbetweenthe
feminine of the Latin word niger, referring to keels, indicating different sources of nest
nearly entirely black body of this species. material. The comb consists of 19 cells, which
REMARKS: This species is most similar to arehexagonalincrosssection.Thecellsinthe
E.scitula,butthesecondmetasomaltergumis central part of the comb are 6–7 mm in
more strongly convex (figs. 4–6) and the diameter (5 side-to-side distance) at opening
female mesoscutum is less convex than in E. and 13–17 mm long; peripheral cells are
scitula; furthermore, E. nigra is also easily somewhat larger (7–8 mm in diameter) and
distinguished from E. scitula by the lack of shorter (9–14 mm in length). The thickness of
bright spots on the clypeus, upper part of the the cell walls is 0.24 mm on average (0.12–
mesopleuron,scutellum,metanotum,andpro- 0.38 mm, N 510). The nest carton is brittle,
podeum. made from small chips mixed with a small
The specific name nigra is van der Vecht’s amount of (possibly salivary) secretion. The
manuscript name. van der Vecht undoubtedly part of the main substrate wire hung from the
distributed to various institutions/museums eaves is thinly covered with a single layer of
the specimens with type labels on which the cartonsheetforabout70 mmfromthebaseof
name appears. We have not yet located the the nest. The other substrate wire tied to the
specimen with his holotype label. mainwireisalsocoveredbyacartonsheet;the
DESCRIPTION OF NESTS: Two nests were coveringstartsabout10 mmfromthenestbase
collected in Tam Dao, northern Viet Nam, and extends for about 100 mm toward the
on9March2005.Oneofthem(nestA,fig. 13) eaves; the part nearthe nest is tubelike (about
was about 2.5 m from the ground, at the 50 mmlongandabout4 mmindiameter),and
junction of two thin (0.5–0.8 mm thick) wires the both ends taper and are closed but it also
hanging from the eaves of a house; the main has an irregularly shaped hole made by in-
substrate wire penetrates the basal part of the completeenclosurenearthemidlength.
nest, with afree end hanging beyond thenest; Theothernest(nestB,fig. 14)wasattached
theotherwireistiedtothemainwire,without to a thin (1.5–1.8 mm thick) twig protruding
a free hanging part. As in all known from a concrete basement, at about 2 m from
Eustenogaster species, the nest has an in- the ground. The nest structure is basically the
verted-flask-shaped envelope. As inall species same as nest A, but the envelope might not
except E. calyptodoma, the envelope is made have been completed and almost lacks in an
by extending the outer walls of peripheral apical tubular extension; the cells are very
cells,andthusitsbasalportioniscomprisedof shallow (3–7 mm long), possibly due to the
the outer walls of peripheral cells. The apical, cell wall having been shaved off at adult
tubular part of the envelope is about 40 mm emergence. The nest color is lighter than in
long and about 10 mm in diameter and is nestA,withthestripesintheenvelopevarying
mesh-walled, with a broad horizontal flange frompalebrowntodarkgrayishbrown.There
near the apex. The upper part of the envelope are 15 cells. Mean thickness of cell walls is
isornamentedwiththree‘‘wings’’thatproject 0.43 mm (0.13–0.72 mm, N 510).
outward,twoofwhichareclosertoeachother WASPSINTHENESTS: NestAhadfourdead
than to the last. The outer surface of the pupaeinthecentralcellsandnoadultsandno
envelopeisfurnishedwithseveralkeels,which live brood. Two of the four dead pupae were
rundowncontinuously,slightlyspirally,tothe so badly molded that we were unable to
apicalflange.Thecolorofthenestisbasically identify their sexes; the other two were a male
brown, with transverse stripes (0.5–1.3 mm and a female.
r
terguminlateralview.8.EustenogasternigraSaitoandNguyen,n.sp.,malemandible.9–12.Eustenogaster
nigraSaitoandNguyen,n.sp.,maleparatype,genitalia.9.Paramereandvolsellaininteriorlateralview.10.
Volsella in interiorlateral view.11. Aedeagusin lateralview. 12.Aedeagus in ventral view.
6 AMERICAN MUSEUMNOVITATES NO. 3534
Figs.13,14. NestsofEustenogasternigraSaitoandNguyen,n.sp.,collectedinTamDao,northernViet
Nam, on9 March 2005.13. Nest A.14. NestB. Scale barsare10mm.
In nest B, 11 adult wasps (6 females and 5 and Khuat, 2004; Nguyen et al., 2006).
males) and no immatures (pupae, larvae, or Eustenogaster hauxwellii seems to be re-
eggs)werepresent.Whenwecollectedthenest stricted to south of the Hai Van Mountains
intoaplasticbag,wetreateditratherroughly, on the border between Thua Thien Hue
but the wasps came out from the nest only and Quang Nam Provinces. On the other
after the nest was kept in the plastic bag for hand, E. scitula extends its distribution
more than 10 minutes. All six females were northward beyond the Hai Van Mountains,
dissected for ovaries and spermathecae under with the northernmost record being from
a stereoscopic dissecting microscope. The Quang Binh Province (fig. 15). Eustenogaster
spermathecae were removed and mounted on nigra is farther north and its southern-
a glass slide in water and examined for most record is from Ha Tinh Province,
presence or absence of sperm under a com- adjacent to Quang Binh Province. If E. nigra
pound light microscope. All six females had and E. scitula do not co-occur and if they are
nearly no fat in their metasoma, undeveloped closely related, this segregated distribution
ovaries,andemptyspermathecae(5uninsem- pattern may imply speciation by a vicariance
inated). event. However, there is no distinct barrier at
least currently between their distribution
DISCUSSION ranges.
DISTRIBUTION
NEST ARCHITECTURE
Besides E. nigra, two Eustenogaster species,
E. hauxwellii (Bingham, 1894) and E. scitula, The nests of Eustenogaster so far known
are recorded from Viet Nam (Nguyen have basically the same structure, except for
2006 SAITO ETAL.: NEW HOVERWASP 7
Fig. 15. Map ofViet Nam showingdistribution recordsof Eustenogaster species.
the envelope, which can be divided into the roots, and thin wires. The first cell (or
followingtwotypes:(1)Envelopearisingfrom sometimes the few first cells) is attached by
the cell wall: the nest with this type of bases to the substrate and further cells are
envelope is usually attached to free-hanging added laterally to the wall of preexisting
fibers, such as plant shoots, free-hanging cell(s). The envelope is made by extending
8 AMERICAN MUSEUMNOVITATES NO. 3534
theouterwallsoftheperipheralcells,andthus and bilaterally symmetric wings at the tip of
it forms a structure continuous to the comb. thetubularentranceseemtoincreasetheeffect
Thistypeofenvelopeisoftencalleda‘‘pseud- ofanesttomimicdeadplantleaves.However,
envelope’’ (Ohgushi et al., 1983; Turillazzi, as discussed by Wenzel and Carpenter (1994)
1991), which implies that the structures are on‘‘crypticnests’’intheNeotropicalpolistine
not true envelopes. Wenzel (1991: 499) dis- tribe Epiponini, the evolutionary process of
agreed with this idea, mentioning ‘‘All enve- thosetraitsbecomingincorporatedintoacryp-
lopes may have been originally derived from tic nest should be discussed with reference to
extensions of the cell wall.’’ Most a robust phylogeny among the taxa con-
Eustenogaster species make nests with this cerned, namely a species-level phylogeny in
type of envelope (see review by Turillazzi, the case of Eustenogaster.
1991). (2) Envelope arising from the substrate In the Stenogastrinae, two types of ant
and independent of the comb: Eustenogaster guard are known. One is an inverted bowl or
calyptodoma (Sakagami and Yoshikawa, an elongate bell made from carton (the same
1968) makes nests on a horizontal or vertical material as the nest) on the threadlike nest
flat surface, where the envelope isconstructed substrate at some distance from the nest,
directlyonthesubstrate,attachingtoitbythe found in Metischnogaster cilipennis (Smith,
base; thus, the envelope is a structure in- 1857), M. drewseni (de Saussure, 1857), and
dependent of the comb. All cells are initiated some Parischnogaster species (reviewed by
directly on (5 attached by their bases to) the Ohgushi et al., 1990). The other is a glob of
horizontalsurface(SakagamiandYoshikawa, secretion from the adult’s Dufour’s gland
1968) or the first cells are attached by their (Sledge et al., 2000), also attached to the
sides to a vertical substrate and further cells threadlike nest substrate at some distance
are added laterally (Hansell, 1987b). Ohgushi from the nest. Such secretion ant guards
etal.(1986)describednestsofEustenogastersp. are known in several species of Parischno-
from Sumatra,inwhich theenvelope basically gaster, rarely in Eustenogaster (review by
arises from the cell walls but is sometimes Ohgushi et al., 1990; see also Carpenter,
partially constructed directly on the substrate 1988), in Liostenogaster tutua Turillazzi, 1999
andisthusindependentofthecomb. (Turillazzi, 1999), and possibly in Stenogaster
ThetwonestsofE.nigraweexaminedhave fulgipennis Gue´rin-Me´neville, 1831 (van der
basicallythestructureofthefirsttype.Among Vecht, 1975). The covering of the substrate
the nests of this type, the E. nigra nest most wire with carton in nest A of E. nigra,
closelyresemblesthatofE.fraterna(Bingham, especially the tubelike portion with the hole
1987) described by Ohgushi et al. (1990) and atmidlength(fig. 13),couldfunctionasanant
the nest of E. eximia (Bingham, 1890) de- guard, but it may also be that the structure
scribed by Krombein (1976, 1991) in having absorbs rain drops running down the wire.
distinct keels and protuberances on the outer Lack of such a covering on the substrate in
surface of the envelope and the apical end of nest B may be because the structure of the
the tubular entrance modified into bilateral substrate, having many branches before the
extensions. The nest of E. nigra differs from wasp nest, functions as an ant guard, or
that of the other two species by the entrance because the nesting site was more or less
tube being distinctly longer (short before protected from rain.
apical extension in the latter two species) and The observation that nest B had produced
the keels on the outer surface of the envelope adults that were still in the course of over-
running slightlyspirally (nearlystraight inthe wintering strongly suggests that E. nigra uses
latter two species). The bilateral extension at a nest for more than one season or more than
the apical end of entrance tube in E. fraterna one generation. Re-use of a nest by
seems to be much less developed than in E. Eustenogaster wasps seems to be commonly
nigra,whereasthatofE.eximiaisoccasionally practiced by succeeding on the natal nest,
much elongated. entering a vacant nest, or by usurpation;
The developed earlike processes, keels on a female could thereby secure the place for
the envelope surface, meshed entrance tube, her brood rearing with a much reduced cost
2006 SAITO ETAL.: NEW HOVERWASP 9
forbuilding(Hansell,1987b).Re-useofanest (no. 18 5900) to F.S. N.L. thanks D.L.
is not common in the social wasps, possibly Khuat, X.L. Truong, H.T. Ta, and V.T.
because of a hygienic problem when the Hoang for help with collecting specimens.
meconia ejected by postfeeding larvae after
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