JHR 97: 531-539 (2024) oe, JOURNAL OF Seewtertensen ern

doi: 10.3897/jhr.97. 123966 RESEARCH ARTICLE () I Tymenopter a
9

https://jhr.pensoft.net The Imarational Sociery of Hymenoperiss, RESEARCH

Distribution of the sand wasp Bicyrtes variegatus
(Oliver, 1789) (Hymenoptera, Crabronidae) in the
Galapagos Islands, with notes on its ecology

Andrea C. Roman!”, Patricio Picébn-Renterfa”?, Charlotte E. Causton!,
Lenyn Betancourt-Cargua', Catherine Frey’, Henri W. Herrera?

| Charles Darwin Research Station, Charles Darwin Foundation, Santa Cruz, Galdpagos, Ecuador
2 Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Laboratorio de Sistemdtica y Biologia
Comparada de Insectos, Bogotd, Colombia 3 Entomology Museum, Facultad de Recursos Naturales. Escuela
Superior Politécnica de Chimborazo. Panamericana sur Km 142, Riobamba, Ecuador

Corresponding author: Andrea C. Roman (andrea.carvajal@fcdarwin.org.ec)

Academic editor: Christopher K. Starr | Received 25 March 2024 | Accepted 1 July 2024 | Published 10 July 2024
https://zoobank. org/83299778-E57B-490E-A LDF-82C17CC11D99

Citation: Roman AC, Picén-Renterfa PR Causton CE, Betancourt-Cargua L, Frey C, Herrera HW (2024) Distribution of the
sand wasp Bicyrtes variegatus (Oliver, 1789) (Hymenoptera, Crabronidae) in the Galapagos Islands, with notes on its ecology.
Journal of Hymenoptera Research 97: 531-539. https://doi.org/10.3897/jhr.97.123966

Abstract

Very little is known about the sand wasp, Bicyrtes variegatus, in the Galapagos archipelago. In this study,
we compiled information from surveys, museum collections and the literature to better understand its
distribution and ecology. We found records of B. variegatus on seven islands with the earliest known re-
cord from 1964, from San Cristdbal Island. Wasps have been collected in the littoral, arid, transition and
humid vegetation zones and have been reported visiting endemic, native, cryptogenic, and introduced
plants. Sun dances and a tight cluster of wasps, similar in form to what others have called “mating balls”,

were observed in the hot season.

Keywords

Bembicinae, flower visitors, mating, sand wasp

Copyright Andrea C. Roman et al. 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.

532 Andrea C. Roman et al. / Journal of Hymenoptera Research 97: 531-539 (2024)

Introduction

Despite the biological importance of the Galapagos archipelago, there are still groups
of insects about which little is known (Causton et al. 2006). Recently, researchers have
increased efforts to study the Hymenoptera of the Galapagos Islands, resulting in the
discovery of new taxa as well as new locality records for species that were already regis-
tered (Fernandez et al. 2018; Herrera et al. 2024). One of these is the sand wasp Bicyrtes
variegatus (Oliver, 1789) (Crabronidae). Species of this genus are recognized by the
scarlike middle ocellus, papal formula (6-4), and lateral projections of the propodeum
(Bohart 1996). Of the 27 described species, 16 are found in South America, eight in
North America, and the remaining species in Panama (Bohart and Menke 1976).

Species from this genus are solitary wasps, feeding mainly on Hemiptera of the
families Alydidae, Coreidae, Cydnidae, Lygaeidae, Pentatomidae, Pyrrhocoridae,
Reduviidae, Rhopalidae, and Scutelleridae, as well as Diptera (Eberhard 1974; Evans
and O’Neill 2007). Evans and O’Neill (2007) identified three methods of provision-
ing nests (mass, delayed, and progressive), depending on the timing and duration of
provisioning a cell, and found that the number of cells per nest ranges from one to five,
with 3—24 prey in a cell. These authors also report nesting-site fidelity, in which wasps
remain in the same place each year. Males display a variety of strategies for attracting
mates including “sun dances”.

Bicyrtes variegatus is easily recognized by the yellow band across the anterior part of
the scutellum (Bohart 1996) and has been reported from Texas to Argentina (Bohart
and Menke 1976; Genise 1979; Amarante 2002; Buys and Trad 2021). Nesting ag-
gregations have also been reported from Santa Cruz Island in the Galapagos (Larsson
1992). Throughout its range, this species is mainly found in coastal habitats, with
observations of females digging nests on the beach (Evans 1976). B. variegatus is con-
sidered a mass provisioner, laying an egg on the prey that it places inside a cell in the
ground nest and then closing the cell. This behavior has been associated with prey scar-
city or poor weather conditions rather than with a particular habitat condition (Evans
and O’Neill 2007). Once the larva matures, a cocoon is made and pupation is initiated.
The cocoon was described by Buys and Trad (2021) as an ovoid and granulose struc-
ture with a length of 9-10 mm. B. variegatus is reported to build nests in the same area
each year, with only the males seeking mates (Evans 1974; Evans and O’Neill 2007).

Very little is known about B. variegatus in the Galapagos Islands. In this study, we
compiled information from surveys, museum collections, and literature to better un-
derstand the distribution and ecology of this species in the archipelago.

Methods

We reviewed specimens deposited in the Invertebrate Collection of the Charles Dar-
win Research Station (ICCDRS) up to August 2023 and specimens from the Califor-
nia Academy of Science (CAS) online database (Monarch 2023). John Heraty kindly

Bicyrtes variegatus in Galapagos 533

provided information on specimens at the University of California, Riverside (UCRC)
and the Canadian Museum of Nature (CMNC), collected during the Peck expedi-
tions (Peck, 2001). The ecological information on the labels or in the databases (e.g.,
interaction with plants) was also extracted. In addition, we conducted a search of the
literature. Lastly, we checked material collected in the project “Study of invasive species
and diversity of terrestrial invertebrates in the Galapagos Islands” within the frame-
work of the ESPOCH agreement with the Charles Darwin Foundation’. In this, a
structured method was used (Longino and Colwell 1997) to inventory Hymenoptera
archipelago-wide in 2018, 2019, and 2021. Sampling was carried out in the littoral,
dry, transition, and humid zones (Jackson 1993; Tye and Francisco-Ortega 2011) on
the islands of Espanola, Fernandina, Floreana, Genovesa, Isabela (Alcedo, Darwin, and
Sierra Negra volcanoes), Marchena, Pinta, San Cristébal, Santa Cruz, and Santiago
using three methods: i) Malaise traps (three traps per zone separated by a minimum of
50 m along the altitudinal transect and set out for eight days), ii) yellow trays (three
trays per zone separated by a minimum of 5 m and placed around each Malaise trap
for three days), and iii) sweep-net collections (four runs of 50 double passes and free
capture in each zone).

Results and discussion

Distribution

The earliest record of B. variegatus that we found was from 1964 collected by R. L.
Usinger on San Cristdébal Island and deposited at CAS. To date, B. variegatus has been
reported from islands currently inhabited by humans (Floreana, Isabela, San Cristébal
and Santa Cruz), inhabited by humans in the past (Santiago; Lundh 2001), and from
uninhabited islands (Espanola and Fernandina). In the material we examined we found
records of B. variegatus from the littoral, arid, and transition vegetation zones (Tye and
Francisco-Ortega 2011), but notably also from the more humid areas (e.g. Bellavista
on Santa Cruz Island and Cerro Pajas on Floreana Island). The highest elevations it
was recorded at was 400 m.a.s.l. NE of Cabo Hammond on Fernandina Island and
335 m.a.s.l. at Cerro Pajas on Floreana Island (Fig. 1). Collection localities include:
Espanola: Gardner Bay; Fernandina: NE of Cabo Hammond; Floreana: Cerro Pajas,
Finca Cruz, Mina de Granillo Negro, Playa Negra, Post Office Bay; Isabela: Bosque de
los Nifos and Puerto Villamil; San Cristébal: Camino a las Negritas, Camino a Tijeras,
El Chino, Puerto Baquerizo Moreno, Puerto Chino; Santa Cruz: Bellavista, Charles
Darwin Research Station (CDRS), Mina Granillo Negro, Playa el Garrapatero, Tor-
tuga Bay. In addition to these records, B. variegatus, incorrectly named Bycertis, is
reported in the literature from Santiago (Traveset et al. 2013) ata site later identified as
Puerto Egas (H.W.H.). No specimens were collected in the archipelago-wide surveys
conducted by ESPOCH between 2018 and 2021, which included setting out traps in
the littoral-arid zones, the preferred habitat of these wasps.

534 Andrea C. Roman et al. / Journal of Hymenoptera Research 97: 531-539 (2024)

N

A

1:1.250,000

4
wy

Marchena Genovesa

Bicyrtes variegatus

& Previous records
@ New records

San Cristobal

| High Zone
Humid Zone :
Transition Zone % a Espanola

Arid Zone = (|
ME Coastal Zone ao)

Figure |. Distribution of Bicyrtes variegatus in the Galapagos archipelago.

How B. variegatus reached the archipelago is unknown. Little information is avail-
able in the literature about how B. variegatus can disperse over long distances, includ-
ing how it has dispersed to new islands (e.g. Bohart and Menke 1976). It is possible
that it arrived in the Galapagos in sand and soil imported for the construction of mili-
tary posts during World War II (US 1947; Peck et al. 1998). These wasps are known
to build their nests in sandy material (Evans and O’Neill 2007) and have been found
in sand used in construction (Eberhard 1974). The cocoons of B. variegatus are made
of soil particles bound together with silk, and the prepupae of some Bicyrtes species
can remain dormant for extended periods (Martins et al. 1998; Buys and Trad 2021).

Interaction with plants and mating behavior

Bicyrtes are known to visit flowering plants in other parts of the world (Porter 1980;
Freitas et al. 2001). Plants are visited to obtain carbohydrates, but are also used for
hunting prey and resting (Martins et al. 1998; Evans and O’Neill 2007). In the
Galapagos, multiple studies have documented Hymenoptera associated with plants
(e.g. McMullen 1986, 1993; Linsley and Usinger 1966; Peck et al. 1998), but did

not report Bicyrtes visiting flowers. However, Bicyrtes sp. was recorded in the literature

Bicyrtes variegatus in Galapagos D9)

(Boada 2005; Traveset et al. 2013; Hervias-Parejo et al. 2018), or on labels, visiting
the following plants: Bidens pilosa (cryptogenic), Croton scouleri (endemic), Galactia
striata (native), Heliotropium angiospermum (native), Lantana camara (introduced),
Mollugo flavescens (endemic), Scalesia cordata (endemic), Scalesia gordilloi (endemic),
Vachellia rorudiana (native), Vallesia glabra (native), Varronia leucophlyctis (endemic),
and Waltheria ovata (native).

Other than plant visits, little is known about the behavior of B. variegatus in
Galapagos. In February 2022, we observed hundreds of Bicyrtes wasps flying near
Garrapatero beach on Santa Cruz Island. The wasps were close to the sand and formed
ageregations akin to mass swarms. These are aggregations of males in the sites where
virgin females emerge (Evans and O’Neill 2007). Such “sun dances” have been docu-
mented as a mating strategy of sand wasps (Bembicinae) (O’Neill 2008). We also saw
a tight cluster of wasps similar in form to what others have called “mating balls” (Evans
and O’Neill 2007), although we did not observe mating (Fig. 2). This behavior was
only observed in February during the hot rainy season. In the following months, only
a few individuals were observed flying over the ground surface.

Figure 2. “Mating ball” of Bicyrtes variegatus on the sand. Credit for illustration: José Falcén Reiban.

536 Andrea C. Roman et al. / Journal of Hymenoptera Research 97: 531-539 (2024)

Material examined

Ecuapor, Galapagos — Espanola ¢ 334, 599; Gardner Bay; 1.35111°S, 89.662222°W;
0-20 m; 27 Apr. 1991; J. Heraty leg.; arid zone; swp; H91-006; UCRC: UCR-
CENT00467383-00467390 — Fernandina ¢ 1; 10 Km N[orth]e[ast] Cabo Hammond;
0.419166°S, 91.581111°W; 400 m; 9 May. 1991; J. Heraty leg.; transition [zone]; H91-
027; UCRC: UCRCENT00467381 — Floreana * 1; Mina de Granillo Negro; 1.281212°S,
90.467887°W; 169 m; 8 May. 2019; A. Acurio, L. Betancourt leg.; ICCDRS 42756 1;
3Km Efast] Playa Negra; 120 m; 21-28 Mar.1989; S. Peck and B.J. Sinclair leg.; Malaise
trap; (89-140); ICCDRS 48230 ¢ 1; Post Office Bay; 30 Abr. 1983; Y. Lubin Leg.; ICCDRS
54389 © 1; Between beach & lagoon, E. end Post Office Bay; 1.235389°S, 90.451361°W;
14 Feb. 1967; Ira L. Wiggins leg; CASENT 8556863 * 2 99; Cerro Pajas; 1.293611°S,
90.457222°W; 335 m; 27 Mar.-22 Apr. 1996; S. Peck leg.; Forest edge; Malaise; 96-54;
CMNC: UCRCENT00467399, 00467400 * 299; Finca Cruz; 130 m, 16-22 Apr.
1996; S. Peck leg.; arid zone forest; Malaise; 96-107; CMNC: UCRCENT00467396,
00467397 * 62 9; Finca Cruz; 130 m; 27 Mar.-16 Apr 1996; S. Peck leg.; arid zone for-
est; Malaise; 96-58; CMNC: UCRCENT00467392-00467395 — Isabela ¢ 1; P[uer]to
Villamil; 7 Mar. 1989; B.J. Sinclair leg.; littoral; sweeping sand beach; ICCDRS 48225
1; Bosque de Nifios; 20 Mar. 1995; P. Delgado leg; Licking on Scalesia cordata; 11:51; IC-
CDRS 48223 — San Cristdébal ¢ 1; Camino a las Negritas; 31 Mar. 2002; R. Boada leg.;
S. gordilloi (leaves); ICCDRS 48221 ¢ 1; Camino a las Negritas, 31 Mar. 2002; R. Boada
leg.; S. gordilloi (flowers); ICCDRS 48228 ¢ 1; [Puerto] Baquerizo; 10 m; 11—23 Feb.
1989; S. Peck leg.; Malaise trap; (89-47); ICCDRS 48224 ¢ 1; El Chino, Z[one] Agricola;
13 Apr. 2003; P. Lincango, A. Mieles leg.; Manual collection (Beach); ICCDRS 48226
1; P[uer]to Baquerizo [Moreno]; 12 Feb. 2007; M. Flores leg.; Manual collection (in the
car); ICCDRS 48227 ¢ 1; Puerto Chino; 26 Feb. 2011; M. Noguera leg.; Manual collec-
tion; on Lantana camara; #SC-544; ICCDRS 54393 ¢ 1; Camino a Tijeras; 2 Mar. 2010;
R. Castro, S. Chamorro leg.; Manual collection; on Croton scouleri; ICCDRS 54390 ° 3;
Progresso (trail to); 0.846271°S, 89.441822°W; 23 Feb. 1964; R.L. Usinger leg.; CASENT
8556860, 8556861, 8556862 — Santa Cruz * 1; C[harles] D[arwin] R[esearch] S[tation];
5 Mar. 1996; C. Blanton leg.; ICCDRS 48222 © 1; Charles Darwin Research Station;
13 Feb. 2007; A. Muth leg.; Malaise; ICCDRS 48229 ¢ 1; Bellavista; 0.6989167°S,
90.3207222°W; 199 m; 12 Feb. 2020; C.R. Gil Jaramillo leg.; Entomological net; IC-
CDRS 46547 ¢ 1; Mina Granillo [Negro]; 8 May. 2011; S. Chamorro leg.; Manual collec-
tion; on Bidens pilosa; ICCDRS 54391 © 1; Tortuga Bay; 23 Feb. 2010; S. Chamorro leg.;
Manual collection; on Waltheria ovata; 1CCDRS 54392 ¢ 2; Playa el Garrapatero; 16 Mar.
2022; A. Mieles Leg; Entomological net; ICCDRS 53891, 53892.

Author contributions

Conceptualization: ACR, PPR. Data curation and formal analysis: ACR, PPR, LBC.
Supervision: HWH, CEC. Writing — original draft: ACR, PPR, CEC. Writing — review
and editing: ACR; ‘CEC, PPR, LBC: CE HWH.

Bicyrtes variegatus in Galapagos 537

Acknowledgments

The Galapagos National Park Directorate (DPNG) and the Ministry of Environ-
ment (MAATE), provided the permits for the annual operation of the Galapagos
Natural History Collections (NPMAATE-MCMEVS-2023-075) under which this
project was conducted. The Study of invasive species and diversity of terrestrial in-
vertebrates in the Galapagos Islands within the framework of the Escuela Superior
Politécnica de Chimborazo (ESPOCH) and Charles Darwin Foundation (CDF)
agreement was conducted with permits from DPNG: PC-43-18, PC-44-19, PC-
43-20, PC-39-21. We thank John Heraty for providing us with new island records
and observations on the manuscript and Kevin O’Neill for his insightful comments,
which improved the manuscript. We also thank Alejandro Mieles for collecting the
specimens from the Garrapatero beach and José Falcén Reiban for the illustration of
the “mating ball” of Bicyrtes variegatus. Funding support for the ICCDRS collection
comes from Gordon and Betty Moore Foundation, Lindblad Expeditions -National
Geographic, and Ecoaventura. This publication is contribution number 2627 of the
Charles Darwin Foundation for the Galapagos Islands, and number 009 under the
agreement ESPOCH-CDFE.

Open Access funding enabled by the project “Study of invasive species and diver-
sity of terrestrial invertebrates in the Galapagos Islands”, within the framework of the
agreement between the Escuela Superior Politécnica de Chimborazo and the Charles
Darwin Foundation.

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