JHR 96: 1031-1044 (2023) yaa JOURNAL OF = 4?eerteewed open-access Jourel

doi: 10.3897/jhr.96. 103379 RESEARCH ARTICLE () Hymenopter a
Q

https://jhr.pensoft.net Te Inarasional Society of Hymenopeeriss, RESEARCH

A new species of social wasp from Madagascar with an
inverted nest architecture (Hymenoptera, Vespidae)

Ozren Polasek!??, Len de Beer*

| Croatian Science Foundation, Zagreb, Croatia 2 University of Split, School of Medicine, Split, Croatia
3 Algebra University College, Zagreb, Croatia 4 Eden Reforestation Projects, Antananarivo, Madagascar

Corresponding author: Ozren Polasek (opolasek@gmail.com)

Academiceditor: Volker Lohrmann | Received 12 March 2023 | Accepted 27 October 2023 | Published 29 November 2023
https://zoobank. org/88FF9B1F-D65E-4F08-B7ED-7A5D40530176

Citation: Polasek O, de Beer L (2023) A new species of social wasp from Madagascar with an inverted nest architecture
(Hymenoptera, Vespidae). Journal of Hymenoptera Research 96: 1031-1044. https://doi.org/10.3897/jhr.96.103379

Abstract

Ropalidia jemmae sp. nov. is described from the protected Ankafobe evergreen forest in central Madagas-
car. This species is characterized by a variable black and green body colour pattern and a unique nest archi-
tecture within the genus Ropalidia. The nests of this species have an inverted cell opening orientation that

exposes the cell bottoms outwardly, mimics the tree bark, and provides excellent visual nest concealment.

Keywords

Nest, new species, protected areas, social wasp, Vespidae

Introduction

Social insects occasionally have very welcome features that supplement the taxonomic
work. In addition to their morphology, some species have unique nest architecture.
This reflects their life cycle, and adaptive mechanisms that can contribute to specia-
tion (Invernizzi and Ruxton 2019). Numerous evolutionary effects can be reflected
in the nest architecture and nesting behaviour (Jeanne 1975), which prevents cata-
strophic brood loss (Furuichi and Kasuya 2015). Interestingly, the nesting architecture
in vespid wasps is considered to have evolved predominantly in response to another
group of social insects, the ants (Richards 1971; Jeanne 1975). Stalked nests, the outer

Copyright Ozren PolaSek & Len de Beer. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

1032 Ozren PolaSek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

envelope or adding the pulp to larva cocoon tops were developed to repel or reduce
invertebrate access to the brood (Jeanne 1975; Furuichi and Kasuya 2015). In contrast
to adaptations protecting against invertebrate predators, visually masking or conceal-
ing nests in the envelopes or tree hollows are believed to have evolved due to vertebrate
nest predation, primarily driven by visual cues (Jeanne 1975).

Among social wasps, the most diverse nest architecture was reported in the genus
Ropalidia (Kojima 1982; Spradbery and Kojima 1989; Wenzel 1998). Members of
the former subgenus Jcarielia, present in Asia and Australia, build enveloped nests
with complex structures (Kojima 1982). On the other hand, most species in this ge-
nus build gymnodomous stelocyttarous nests, characterized by the stalked nest with-
out any envelope (Jeanne 1975; Wenzel 1998). Most African mainland species build
simple discoid nests (Polasek et al. in press), while Malagasy Ropalidia exhibit much
greater nest architecture diversity (Vesey-Fitzgerald 1950; Blommers 2012).

All 43 Malagasy Ropalidia species are endemic (Carpenter and Madl 2009; Blom-
mers 2012). Despite substantial obstacles in their separation due to the lack of a taxo-
nomic revision, some species were shown to have unique nest architecture, which
provides a valuable supplement to species determination (Blommers 2012). Nota-
bly, some Malagasy species build visually concealed nests directly on the tree trunks
(Vesey-Fitzgerald 1950), a feature described in only one African mainland species
(Polasek et al. 2022). In addition, R. cocoscola Blommers was reported to build entirely
concealed nests within the tree trunk cavities, while R. merina Blommers (previously
considered as R. formosa de Saussure) and R. favulorum Blommers create large nest-
ing colonies of hundreds of nests that provide an additional layer of security (Wenzel
1987; Blommers 2012).

The aim of this study was to identify and describe a taxon that was recently observed
in a protected area of the Ankafobe forest. The Tampoketsa de Ankazobe, including the
Ambohitantely forest, is one of the last and the largest remaining fragmented forests on
the highlands of Madagascar, a mosaic of forest and grassland. Ambohitantely forest
consists of about around 80 sections of the subhumid, high plateau evergreen forest at
an altitude between 1,300 and 1,650 metres. With about 1800 hectares of surface area,
it harbours many endemic and critically endangered species of insects (Wesener et al.
2010; Wiorek et al. 2021; Moravec and Trzna 2022), frogs and mammals (Stephenson
et al. 1994; Barata et al. 2022), requiring careful management, protection and restora-
tion efforts.

Materials and methods

This study was based on the field observation of the wasps on a nest in the Ankafobe
forest in October and November 2022. ‘The initial observation prompted a wider-
scale search for the corresponding specimen in the accessible entomological col-
lections. In total, 18 collections were examined, including the American Museum

of Natural History, New York, US [AMNH], Centre for Biodiversity Genomics,

New social wasp with inverted nest 1033

Guelph, Ontario, Canada [CBCG], Centrum ftir Naturkunde, Zoological Muse-
um, Universitat Hamburg, Germany [CeNaK], Entomological Collection, ETH
Zurich, Switzerland [ETH], Hungarian Natural History Museum, Budapest, Hun-
gary [HNHM], Museo di Storia Naturale di Venezia, Italy [MSNV], Muséum
d Histoire Naturelle, Geneva, Switzerland [MHNG], Museum ftir Naturkunde,
Berlin, Germany [MFNB], National Museum of Natural History, Naturalis, Lei-
den, the Netherlands [NNM], Natural History Museum London, UK [NHM],
National Museum Kenya, Nairobi, Kenya [NMK], National Museums Scotland,
Edinburgh, Scotland [NMS], Oberdsterreichisches Landesmuseum, Linz, Austria
[OLM], Royal Museum for Central Africa, Tervuren, Belgium [RMCA], Swedish
Museum of Natural History, Stockholm, Sweden [NHRS], Statens Naturhistor-
iske Museum, Copenhagen, Denmark [SNM], Zoologisches Forschungsmuseum
Alexander Koenig, Bonn, Germany [ZFMK] and Zoologische Staatssammlung
Miinchen, Munich, Germany [ZSM].

All specimens were examined under a Leica S9i stereomicroscope and photo-
graphed with an integrated camera. Images were stacked using Helicon Focus version
6.8.0 (Kharkiv, Ukraine). All terga and sterna were denoted as T or S, while the flagel-
lomeres were denoted as AF1—AF10.

Results

Taxonomy

Ropalidia jemmae Polasek & de Beer, sp. nov.
https://zoobank.org/13ADD49A-3E0A-45B6-9889-22AB94EB40D4
Figs 1-5, holotype labels Fig. 6

Material. Holotype: “Ankafobe/Ambohitantely”, 19. Paratypes: the same location as
the holotype, 29 9. The nest from the same series also has an additional label “5. 52/
(RP)”, [nest A] (all in MSNYV).

Observation data. Ankafobe [-18.10492, 47.187227], 27 Oct 2022 and 10 Nov
2022, 229 anda nest [nest B], obs. Len de Beer [https://www.inaturalist.org/observa-
tions/ 141686572, https://www.inaturalist.org/observations/ 140282514].

Diagnosis. This species is characterized by the combination of the following char-
acters: I'l shape streamlined, propodeal excavation impunctate, black basal colour and
variably expressed whitish-green markings on the head and mesosoma, green markings
only on coxa II, widened posterior yellow-green bands on T1 and T2, and overall
longer pilosity. Wing length: 7.9 [7.8-8.0] mm.

Description. Female. Colour. Basal body colour black (Figs 1, 2). Head black,
with green line in apical fourth of clypeus (Fig. 3). Interantennal area and inner orbits
with faint green markings, mandible with basal green spot (Fig. 3). Gena and tem-
pora suffused brown-green (Fig. 1). Antennal scape, pedicel and AF1 green ventrally,

1034 Ozren PolaSek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

flagellum dorsally black, remaining flagellomeres ventrally orange (Fig. 1). Pronotum
suffused green, with stronger green hue close to carina; inferior angle darker; mes-
onotum without green markings (Fig. 2). Suffused green spot high on mesopleuron,
posterior third of tegula green (Figs 1, 2). Scutellum with two green spots; metanotum
with two green spots that have stronger margins than those on scutellum (Fig. 4). Pro-
podeum black (Fig. 4). Coxa I and III black, coxa II with lateral green quadrant (very
faint in one specimen). Femora black with elongated or circular green spot near distal
margin on femora II and III. Tibia with two shades of green; tarsi green, distally light
brown (Figs 1, 2). Wings translucent, with some yellowing anteriorly; nervature yellow
to brown, stigma yellowish and translucent, apical spot faintly developed in anterior
third of marginal cell (Fig. 1). T1 with posterior yellow band that occupies about half
of the total surface, thin connecting green suffused area and a dorsal remaining black
diamond-shaped area (one paratype has mainly black T1, with thin remaining green
band). T2 with posterior widened band, characterized by two larger attached areas
and two remaining spots in basal colour near posterior margin, located close to T2/
S2 suture (Fig. 1; band substantially reduced in one paratype, shown in Fig. 5). This
pattern extends on S2 as short and smaller yellow-green spot, integrated with posterior
band (very reduced in one specimen). Remaining terga and sterna in basal colour or
somewhat brownish.

Head. Head frontally 1.2x as wide as high; clypeus 1.15x as wide as long (Fig. 3).
Clypeal base elongated, lateral margins parallel, juxtamandibular lobe moderately de-
veloped, apex pointy and projecting (Fig. 3). Clypeus surface with basal sculpture and
evenly spaced and well-defined small punctures, obscured by pubescence; punctures
on apex coarser and converting into poorly defined craters (Fig. 3). Frons with dense,
moderately sized and comparatively shallow punctures, vertex behind ocelli with di-
minishing punctures, area close to occipital carina impunctate and shiny. Gena and
tempora with equally sized, but shallower punctures. Gena posteriorly 0.6—0.8x as
wide as eye, mainly parallel to posterior eye margin; occipital carina complete, reaches
mandible. Clypeus and frons covered by dense and straight silvery-yellowish pubes-
cence and yellowish protruding setae, about equally long as forward ocellus diameter
(Fig. 3). Compound eyes setose (Fig. 3). Ocellar triangle equidistant; distance between
lateral ocelli about 0.6x of ocelli-eye distance. Scape equally long as AF1, AF1 equally
long as AF2+3+4, AF2 about 1.4—1.5x as long as wide.

Mesosoma. Mesosoma about 1.8x as long as wide in dorsal view, mesonotum
1.2x as long as wide (Fig. 2). Head wider than pronotal carina width (Fig. 2). Pronotal
carina developed as translucent rim, twice wider dorsally than laterally; entire prono-
tum covered by shallow and very dense punctures separated from each other by their
diameter; inferior pronotal corner with increasingly larger punctures (Fig. 1). Mes-
onotum punctures less dense and shallow, somewhat denser close to scutellum. Meso-
pleuron densely punctate. Metapleuron shiny, with an occasional very weak punctum
that can only be visualized in specimen rotation. Scutellum densely punctate, with
shallower punctures than those on mesonotum. Median scutellar carina developed and
thin, reaches about half of scutellar length. Metanotum dorsally punctate, with shiny

New social wasp with inverted nest 1035

impunctate posterior triangle with small projecting area towards propodeum. Propo-
deal excavation shallow, without strong carina; upper half with cuticular surface, lower

with minute striae; entire propodeal excavation completely impunctate (Fig. 4). Entire
mesosoma covered by silvery-yellowish pubescence and some protruding setae; setae

2M gga. a Abed

Figures 1-6. R. jemmae sp. nov., female, habitus | lateral view 2 dorsal view 3 head in frontal view

4 propodeal excavation 5 distal margin of T2 6 label of the holotype.

1036 Ozren Polasek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

on dorsal side of pronotum silvery, those on mesonotum yellowish. Propodeal excava-
tion covered by longer silvery setae.

Metasoma. 11 pyriform and elongated, with developed dorsal part, but not glob-
ular in shape; posterior sulcus very weak (Figs 1, 2). T1 shallowly punctate, punctures
poorly defined. T2 shiny, with shallow, dense and small punctures, covered by silvery
pubescence and longer silvery-yellowish protruding setae that extend well over lamella
(Fig. 5). [2 somewhat shorter than S2, visible in dorsally oblique lamella (Fig. 4). Re-
maining metasomal segments with weakly developed punctures and pubescence longer
than that on T2 and S2.

Figures 7-12. R. jemmae sp. nov. nest 7 nest B, in situ, lateral view 8 with wasps 9 nest A, cell openings
10 lateral view II outer view (notably, the spot in the middle is the glue that connected the nest to the

underlying cardboard) 12 greater magnification, showing a brittle paper structure.

New social wasp with inverted nest 1037

Males are unknown (two more pre-hatching larvae were recovered from the co-
coons of nest A, macerated and examined, but both were females). Notably, at least
one male was observed on nest B (Fig. 8), with entirely yellow clypeus. However, in the
absence of the specimen, the male remains undescribed.

Similar species. Several Malagasy species have a similar morphology and a gen-
eral colour pattern. In order to provide sufficient support to separating R. jemmae
sp. nov. from previously described species, a partial key is provided here. The key
is designed to separate the four similar species from others (key couplet 1). The key
couplet 2 can be treated as 25b in the previous key to Malagasy species (Giordani
Soika 1991).

Nest. The nest is the single most interesting feature of this species, with unique
architecture in the entire genus Ropalidia. Instead of the cell openings oriented
outwardly, the nests of R. jemmae sp. nov. are inverted, with the cell openings ori-
ented towards the nesting surface and the cell bottoms oriented outwardly (Fig.
7). The loosely built cell bottoms thus correspond to the rugged texture of the tree
bark or the lichen and provide excellent concealment of the nest (Fig. 8). In ad-
dition, the nest is built with greenish-grey material, mostly homogenous, without
streaks of different colours integrated into the cell walls, which are common in
some other species.

The first examined nest, nest A, has a total of 43 cells, arranged in the 7*9 cells
maximum. The nest is somewhat elongated, with six enclosed cells (Fig. 9). The paper
structure is brittle, visible in several collapsed cells (Fig. 9). Approximately 50%-75%
of the cell length overlap between the cell rows (Fig. 10). The stalk of the same nest
is also inverted, suggesting attachment on the cell openings side (Fig. 10). The outer
(bottom-exposed) side of the cells has a very textured surface with numerous arches,
which do not reflect the cell wall structure but are located much more densely, pro-
viding a heavily textured outer surface (Fig. 11). The greater magnification shows no
elongated fibres, but only clumps of heavily masticated and mostly rounded chips
(Fig. 12).

The second, nest B, was only observed in situ (Figs 7, 8). The nest was on the tree
trunk, 1.3 m above the ground. The nest colour and shape resemble the nearby greyish-
green lichen patch (Fig. 7).

In comparison, the nests of R. ranavali are elongated with a pointy tip, suspended
from similarly coloured branches (https://www.inaturalist.org/observations/63 188743,
https://www.inaturalist.org/observations/9173358). The nests of R. venustula and
R. scottiana are unknown.

Etymology. The name is given after Jemma de Beer, who participated in the nest
B discovery.

Distribution. All the examined specimens and observations originated from a nar-
row area (of no more than 16 km of distance) in central Madagascar near Ankafobe, at
the edge of the Ambohitantely forest.

1038 Ozren Polasek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

Partial key to species

1

T1 streamlined and posteriorly rounded, with poorly defined punctures
(a—c). Basal body colour black(ish)-olivaceous, T2 with obligate posterior
green or yellow band on T1 and T2, with additional four attached spots on
T2 that can be small in size (d), intermediate and merged (e) or occupy most
of T2 surface (f). Femora II and II black, sometimes with distal green or yel-
low spot. Mesonotum black. Wing length 7.5-10.3 mm oie eeeeeeeeeeeees 2

» oe

T1 differently shaped, dorsally developed (aa) and distally punctate (bb),
more angulate (cc) or with stronger posterior sulcus (dd). Basal body colour
variable; if there are green or yellow markings on T2 then those are either
just a posterior band (ee), bilateral green-yellow spot near base (ff) or entire
T2 surface green or yellow (gg). Femora II and III variably coloured (if T2
has wider posterior band, then femora are green). Mesonotum variably col-
oured (if T2 has wider posterior band, then mesonotum always with a pair
ole reenslone ina ivalelities) io: .t.rt mJ. mans centers, Nadas UPR SEED. St SRD
Radetich Ate. other species, according to the Giordani Soika (1991) key

New social wasp with inverted nest 1039

Head and mesosoma with numerous green or yellow markings (a). Pronotal
carina twice broader dorsally than laterally, commonly with a fine translucent

rim (b). Base of clypeus with poorly defined punctures (c). Femora II and II
blackawithvdistally cllowsor-ereen spot. (Gd): cess andere Rens ee erate, 3

Clypeus, interantennal area, pronotum, scutellum, mesopleuron and metano-
tum commonly black, at most with olivaceous suffused greenish colouration
(aa). Pronotal carina equally wide dorsally and laterally, without translucent
rim (bb). Base of clypeus with large and well-defined punctures (cc). Femora
II and III at most with a very faint suffused olivaceous spot, commonly en-

tirely black (dd)

Face with numerous yellow markings, clypeus entirely yellow (a). Clypeus
wider apically, pentagonal (b). Eyes asetose (c). Propodeal excavation shiny,
with more developed striae, present in upper half (d); markings on body
yellow-green (e). AF2 as long as wide... R. scottiana (de Saussure)

1040 Ozren Polasek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

— Face mostly black, with lesser amount of greenish-white markings (aa). Cl-
ypeus elongated, with parallel sides (bb). Eyes covered by numerous straight
setae (cc). Propodeal excavation shiny, upper half without punctures or striae
(dd); markings greenish-white (ee). AF2 about 1.4—1.5x as long as wide......
Foi emer ieee ne Ene gia rnc acs er core ain renee ERT EL eed or eet R. jemmae sp. nov.

4 T2 with broad posterior yellow band (a). Distal margin of T2 towards lamella
evenly rounded and straight; lamella with linear digitations, impunctate (b).
5 ane 5, npr myer yon 6-8 maaan 5 Scns, lee 0 0 ioe peel R. ranavali (de Saussure)

New social wasp with inverted nest 1041

— T2 with thin posterior whitish-yellow band with integrated spots (aa). Distal
margin of T2 towards lamella serrated, with translucent rim; lamella coarsely
punctate Dib) vete deem urna dress emesteme noes ie R. venustula (de Saussure)

Discussion

The new Ropalidia species exhibits an interesting case of an inverted nest, which can be
considered an evolutionary reversal. Instead of favouring economically and structurally
stronger ancestral nest type, with regular cell size and shared adjacent walls, this species
evolved a trade-off by favouring visual concealment. Such architecture requires more
founder and worker effort than ancestral nests, which might directly contribute to the low
abundance and scarcity of collected or observed specimens. ‘This kind of concealing of the
nest is unique in Ropalidia, but a similar example of visual concealment is known from the
Neotropical species of Mischocyttarus (Milani et al. 2021; O’ Donnell 2021), suggesting that
visual predation of social wasps nests is a common evolutionary pressure across continents.

The biology of R. jemmae sp. nov. remains unknown, but this is similar to almost
all Malagasy Ropalidia, with only a handful of documented field observations (Vesey-
Fitzgekald 1950; Wenzel 1987; Blommers 2012). Lastly, the declining habitat un-
doubtedly presents another negative factor towards species abundance, suggesting the
need to reverse this trend, propelled by the restoration of the native forests.

The green cuticular colour in Malagasy Vespidae is a strong indicator of the selec-
tive pressure exerted on the nests and the adult wasps. Interestingly, the same feature is
known not only for numerous Ropalidia species, but also for several Belonogaster spe-
cies (Hensen and Blommers 1987), and at least one eumenid species (Giordani Soika

1042 Ozren PolaSek & Len de Beer / Journal of Hymenoptera Research 96: 1031-1044 (2023)

1941). This pattern of local camouflage can be viewed as both colour matching (fit-
ting the background colour) and disruptive colouring (the pattern of green and black
areas on the body that cause visual confusion and break the expected body contour).
It seems that the adaptive radiation in Madagascar has taken more concealed pathway
in vespid wasps, as opposed to bright black-and-yellow species present in temperate
regions or brown- red-black predominant colour in tropical regions.

The conservation efforts in Malagasy forests are of utmost importance in maintain-
ing their biodiversity. The restoration efforts, while present in the area, are hampered
by cultural practices, exploitation of the forests, and invasive species. Additional effort
is warranted, in order to retain the known and unknown biodiversity of this irre-
placeable area. The mosaic of forest and grassland called the Tampoketsa d’ Ankazobe,
which is the only proven habitat for Ropalidia jemmae sp. nov., has the disadvantage
of proximity to Madagascar’s capital city and has been subject to years of exploitation
for wood in an area, where the use of fire regularly during the dry period, has almost
totally decimated what was once a diverse mix of trees and grass.

Protective and restorative efforts focused on the remaining fragments of evergreen
forests in the Tampoketsa, namely Ankafobe and Ambohitantely are of paramount im-
portance in retaining their utterly unique elements of biodiversity that are not found
anywhere else, including in some cases not even anywhere else in Madagascar. Several
NGOs are undertaking restorative reforestation under challenging conditions.

Acknowledgements

A very special thanks goes to all researchers and museum curators, who assisted in
creation of this paper (in alphabetical order): Frederique Bakker (NNM), Gavin Broad
and Andrew Polaszek (NHM), James M Carpenter (AMNH), Michael Greeff and Rod
Eastwood (ETH), Allison Brown and Gergin Blagoev (CBCG), Martin Husemann
and Thure Dalsgaard (CeNaK), Bernard Landry (MHNG), Laban Njoroge (NMk),
Michael Ohl and Lukas Kirschey (MFNB), Ralph S. Peters (ZFMK), Lars Vilhelm-
sen (SNM), Martin Schwarz (OLM), Vladimir Blagoderov (NMS), Olga and Stefan
Schmidt (ZSM), Marco Uliana (MSNV), Didier van den Spiegel and Stéphane Hanot
(RMCA), Hege Vardal (NHRS) and Zoltan Vas (HNHM).

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