European Journal of Taxonomy 211: 1-12

http://dx.doi.org/10.5852/ejt.2016.211

BY

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2016 • Prebus M. & Lubertazzi D.

Research article

urn:lsid:zoobank.org:pub:9A706453-A104-45FB-8B9C-92C8D18723A4

A new species of the ant genus Bothriomyrmex Emery, 1869

(Hymenoptera: Formicidae) from the Caribbean region

Matthew PREBUS 1 * & David LUBERTAZZI 2

’Department of Entomology & Nematology, University of California, Davis, Davis, CA 95616, USA.

2 Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.

^Corresponding author: [email protected]

2 Email: [email protected]

1 um:lsid:zoobank.org:author:lA6494C7-795E-455C-B66F-7F6C32F76584

2 urn:lsid:zoobank.org:author:3C9C4B01-403B-4A38-973E-F86B4B06D494

Abstract. Bothriomyrmex enigmaticus sp. nov. is described from the island of Hispanola based on one

nest collection. This is the first collection of the genus Bothriomyrmex (Emery, 1869) from the Caribbean

region, and the second species to be described from the Americas. While sharing several characters

with B. paradoxus (Dubovikoff & Fongino, 2004) from Costa Rica and Honduras, B. enigmaticus sp.

nov. diverges in several key characters, including palp formula. However, a morphometric comparison

to Palearctic species of the tribe Bothriomyrmecini suggests affinities to B. paradoxus , Chronoxenus

wroughtoni (Forel, 1895) of the eastern Palearctic, and to a lesser extent an undescribed species of

Arnoldius (Dubovikoff, 2005) from Australia and B. corsicus (Santschi,1923) of the western Palearctic.

Keywords. Bothriomyrmecini, Dolichoderinae, Greater Antilles, disjunct distribution. Neotropic.

Prebus M. & Lubertazzi D. 2016. A new species of the ant genus Bothriomyrmex Emery, 1869 (Hymenoptera:

Formicidae) from the Caribbean region. European Journal of Taxonomy 211: 1-12. http://dx.doi.org/10.5852/

eit.2016.211

Introduction

Bothriomyrmex (Emery, 1869), with 23 described species (Bolton, 2014), is a small genus of ants in the

subfamily Dolichoderinae (Forel, 1878). With an unusual distribution, encompassing the Palearctic and

Neotropic regions, Bothriomyrmex has been collected in a broad range of habitats, including grassland,

savannah, lowland tropical moist forest and cloud forest. Nests typically occur in the soil, under rocks

or in rotten wood. While little is known about the diet of many species, Santschi (1919) noted that

Bothriomyrmex preferred a diet of sugar water, and observed a nest of B. crosi (Santschi, 1919) with

nymphs of Tettigometra (Latreille, 1804), suggesting trophobiosis as one source of nutrition. Other

details of the life history of many species remain obscure, but Forel (1873) was the first to observe a

mixed colony of B. meridionalis (Roger, 1863) and Tapinoma erraticum (Latreille, 1798) in Switzerland.

Later, in North Africa, Santschi (1906) observed that newly mated queens of Bothriomyrmex invade

colonies of Tapinoma and usurp their nests by decapitating the queen over the course of several days,

1

European Journal of Taxonomy 211 : 1-12 ( 2016 )

suggesting that at least in some cases Bothriomyrmex is a temporary social parasite. Due to the peculiar

mandibular morphology of all known gynes in this genus, Seifert (2012) asserted that all species of

Bothriomyrmex are temporary social parasites, with nest founding strategies similar to the one observed

by Santschi. A study by Lloyd et al. (1986) suggested that the anal gland secretions of B. syrius (Forel,

1910) gynes facilitate their entrance into host nests, via chemical mimicry of the Tapinoma colony odor.

In 2004, a new species of Bothriomyrmex, B. paradoxus (Dubovikoff & Longino, 2004), was described

from material collected in Costa Rica; further collections were made in mesophyll forest in Honduras

during the 2010 Leaf Litter Arthropods of Meso-America (LLAMA) expedition. Prior to this, this

genus was considered to be geographically restricted to the Palearctic, Oriental and Australian regions.

Recently, Bothriomyrmex was separated into 3 geographically graded genera based on palp formula and

gyne wing venation (see Table 1): Bothriomyrmex s.s., with 23 described species from the Palearctic

region; Chronoxenus (Santschi, 1919), with 6 described species from the Oriental region; and Arnoldius

(Dubovikoff, 2005), with 3 species from the Australian region (Dubovikoff 2005). Bothriomyrmex

paradoxus , with palp formula 4,3 and a closed discoidal cell, is thought to be a member of Bothriomyrmex

s.s.: indeed, a 2010 study in which the molecular phylogeny of the subfamily Dolichoderinae was

estimated provided additional evidence for this argument (Ward et al. 2010, see Fig. 1).

In this article we describe a second species of Bothriomyrmex from the Americas. This species was

found on the the island of Hispanola, marking the first record of this genus in the Caribbean region.

Additionally, we utilize a morphometric dataset developed by Seifert (2012) in an effort to develop a

hypothesis that could explain the disjunct distribution of this genus.

Arnoldius AU01

Chronoxenusjavanus

Bothriomyrmex saundersi

Bothriomyrmex paradoxus

Fig. 1 . Phylogenetic relationships among three closely related genera in the tribe Bothriomyrmecini,

based on Ward et al. (2010).

2

PREBUS M. & LUBERTAZZID., Caribbean Bothriomyrmex

Table 1 . A summary of characters separating three closely related genera in the tribe Bothriomyrmecini,

based on Dubovikoff (2005). *2,3 in Bothriomyrmex enigmaticus sp. nov. (this study).

palp formula

gyne forewing venation

biogeographic region

Bothriomyrmex s.s.

4,3*

closed discoidal cell

Palearctic, Neotropic

Chronoxenus

2,3

closed discoidal cell,

reduced cubital and

medial veins

Oriental

Arnoldius

2,2

open discoidal cell

Australian

Material and methods

Abbreviations of depositories

The material upon which this study is based is located and/or was examined at the following institutions:

CAS

INBio

LACM

MNHNSD

MHNG

MCZ

UCDC

USNM

ZIN

California Academy of Sciences, San Francisco, CA, USA

Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica

Los Angeles County Museum of Natural History, Los Angeles, CA, USA

Museo Nacional de Historia Natural “Prof. Eugenio de Jesus Marcano”, Santo Domingo,

Dominican Republic

Museum d’Histoire Naturelle, Geneva, Switzerland.

Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA

University of California, Davis, Davis, CA, USA

National Museum of Natural History, Washington, DC, USA.

Zoological Institute, Russian Academy of Sciences, Saint Petersburg, Russia

In this study we compare the morphology of Bothriomyrmex paradoxus and B. enigmaticus sp. nov.

Because the holotype of B. paradoxus is a gyne, and the gyne of B. enigmaticus sp. nov. remains

unknown, we examined paratype workers of the former as a basis for comparison. These paratypes

came from the collections of CAS [JTLC000003512] (images examined on www.antweb.org ). MHNG

[CASENTO103277] (images examined on www.antweb.org T and UCDC [JTLC000003511],

The type material of the new species and all imaged specimens are uniquely identified with specimen-level

codes affixed to each pin (e.g. CASENT0636109). Digital color images and measurements were taken

using a Leica MZ 12.5 stereo microscope equipped with a JVC KY-F75 digital camera and Syncroscopy

Auto-Montage software (version 5.0). All images were taken by the authors, except where indicated.

The images and specimen data presented are available online and can be viewed on AntWeb ( www.

antweb.org ). Measurements and indices are presented as minimum and maximum values, expressed

in mm to three decimal places, with arit hm etic means in parentheses. The following measurements are

derived from those used by Seifert (2012) and Dubovikoff & Longino (2004).

Measurements

CL

ClyW

CS

CW

dAN

EL

Cephalic length: maximum distance from the mid-point of the anterior clypeal margin to the

mid-point of the posterior margin of head, measured in full-face view

Maximum clypeal width in full face view

Cephalic size; arithmetic mean of CL and CW

Cephalic width: maximum width of head measured in full face view, excluding the eyes

Minimum distance between the inner margins of the toruli (the ring sclerites surmounting

and enclosing the antennal sockets)

Eye length: maximum diameter of compound eye including all structurally visible ommatidia

3

European Journal of Taxonomy 211: 1-12 (2016)

EW

EYE

F2

IF 2

MGr

MW

PoOc

PH

PrL

SL

WL

irrespective of the pigmentation status, measured in oblique lateral view

Eye width: minimum diameter of compound eye including all structurally visible ommatidia

irrespective of the pigmentation status, measured in oblique lateral view

Eye size: arithmetic mean of EL and EW

Median length of 2 nd funicular segment in dorsal view; take care to measure median length

because of unequal length of the segments’ sides. Use the highest optical resolution available

F2 divided by the maximum width of second funicular segment (excluding pubescence) in

dorsal view

Depth of metanotal groove/depression in profile view; the upper reference line extends

between the highest points of mesonotum and propodeum perpendicular to which depth

measuring is performed. Use of transmitted light ensures the most accuracy

Pronotal width: maximum width of pronotum measured in dorsal view

Postocular distance: distance from the transverse level of posterior eye margin to hind

margin of head, measured medially

Petiole height: ma xim um height of the petiole in profile view, including protruding lobe on

the ventral margin

Propodeal length: distance from most caudal point of posterior propodeum (a dentiform para¬

median structure on both sides of petiolar junction) to hind the margin of the metanotal groove

Scape length: maximum antennal scape length excluding basal condyle and neck

Weber’s length: the diagonal length of the mesosoma in profile from the point at which the

pronotum meets the cervical shield to the posterior basal angle of the metapleuron

The varying degree of inclination of pubescence and pilosity is often of high diagnostic value throughout

a broad spectrum of ant genera. In this context we use the terms “erect”, “suberect”, “subdecumbent”,

“decumbent”, and “appressed” following the usage in Wilson (1955).

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Suborder Apocrita Latreille, 1810

Family Formicidae Latreille, 1809

Subfamily Dolichoderinae Forel, 1878

Tribe Bothriomyrmecini Dubovikoff, 2005

Genus Bothriomyrmex Emery, 1869

Synopsis of Neotropical Bothriomyrmex species:

Bothriomyrmex paradoxus (Dubovikoff & Longino, 2004)

Bothriomyrmex enigmaticus sp. nov.

urn:lsid:zoobank.org:act:A78CBE68-03A8-4A17-A0A7-44C76AC166F8

Figs 2b-e, 3M

Diagnosis

The following character combination distinguishes Bothriomyrmex enigmaticus sp. nov. from

B. paradoxus : palp formula 2,3; medial lobe of clypeus not strongly projecting: flat, or with a broad,

shallow concavity anteriorly. Long, paired setae on medial clypeal lobe separated by 2 /s their length or

more. Eyes large: OI 23.1-24.7. Head box-lilce, with occipital corners narrowly rounded.

4

PREBUS M. & LUBERTAZZID., Caribbean Bothriomyrmex

Etymology

The specific epithet is derived from the ancient Greek “aiviypaiiKO^” meaning “riddle”, and bears

a double entendre: not only is this the second species in this genus with a paradoxical distribution,

this species also presents a puzzling new perspective on the generic classification system of the tribe

B othriomy rme cini.

Type material examined

Holotype worker

DOMINICAN REPUBLIC: Provincia Maria Trinidad Sanchez, 7 km WSW El Factor, 19.29776° N,

69.94977° W ±3 m, 195 m ±5 m, 21 Jul. 2015, disturbed broadleaf moist forest, nest in dead top of live

tree, M.M. Prebus#MMP01990 [MCZ-ENT00035850],

Fig. 2. Clypeal structure, a. Bothriomyrmex paradoxus (Dubovikoff & Longino, 2004) (JTLC000003511).

b. B. enigmaticus sp. nov. (MCZ-ENT00035850); arrows indicate the insertion points of long setae

on anterior clypeal margin, c. Petiole, d. Mouthpart dissection, e.Ventral view of head capsule of

B. enigmaticus sp. nov. (MCZ-ENT00035850). Photo credit for 2a: Michael Branstetter. All scale bars

represent 0.2 mm

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European Journal of Taxonomy 211: 1-12 (2016)

Fig. 3. Holotype worker of Bothriomyrmex enigmaticus sp. nov. (MCZ-ENT00035850). a. Full face

view. b. Profile view. c. Dorsal view.

6

PREBUS M. & LUBERTAZZID., Caribbean Bothriomyrmex

Paratypes

DOMINICAN REPUBLIC: Provincia Maria Trinidad Sanchez, 9 workers, same collection as holotype

[CAS: CASENT0756066; INBio: CASENT0756067; LACM: CASENT0756068; MCZ: MCZ-

ENT00539146, MCZ-ENT00539147; MNHNSD: MNHNSD 18.106; UCDC: CASENT0756071;

USNM: CASENT0756072; ZIN: CASENT0755363],

Worker description

Measurements (n=ll): CL 0.541-0.563 (0.554); ClyW 0.375-0.412 (0.395); CS 0.520-0.542 (0.531);

CW 0.495-0.523 (0.508); dAN 0.181-0.215 (0.197); EL 0.124-0.137 (0.132); EW 0.096-0.109 (0.105);

EYE 0.112-0.122 (0.118); F2W 0.062-0.070 (0.065); IF2 1.000-1.222 (1.117); MGr 0.019-0.030

(0.024); MW 0.318-0.357 (0.343); PoOC 0.251-0.274 (0.260); PH 0.214-0.250 (0.231); PrL 0.264-

0.296 (0.282); SL 0.407-0.434 (0.424); WL 0.572-0.617 (0.600).

Palp formula 2,3; distal maxillary palp segment roughly twice as long as basal (Fig. 2d). Medial hypostoma

absent (Fig. 2e). Masticatory margin of mandible with 6 teeth and 1-2 denticles. Mandibles with short,

curved setae similar to those on the anterior clypeal margin. Clypeus narrow: 0.10-0.11 mm high

medially. Anterior clypeal margin flat, or with a broad, shallow concavity; bearing many short, curved

setae; one long seta on each side of concavity, their bases separated by the length of the setae or more.

Medial lobe of clypeus not strongly projecting. Posterior margin of clypeus even with anterior surfaces

of antennal socket cavities. Antennal scapes short, not reaching the posterior margin of the head in full

face view. Head slightly oblong (CL/CW 1.069-1.108), with lateral margins evenly convex; widest part

of head in full-face view posterior to the compound eyes. Posterior head margin flat, becoming slightly

concave medially; corners of head narrowly rounded, giving the entire head a blocky appearance. Head

with two long setae on frons (longer than the first funicular segment) and two shorter setae on posterior

clypeal margin (shorter than the first funicular segment), otherwise covered uniformly with short, dense,

decumbant pubescence. Eyes large (EYE/CS 0.214-0.229), 8 ommatidia in longest row.

Posteroventral pronotum margin narrowly rounded. Metanotal groove deeply impressed. Propodeum

high and rounded, with declivitous face roughly twice as long as dorsal face in profile; propodeal angle

indistinct. Mesosoma covered uniformly with short, dense, decumbent pubescence.

Fig. 4. Larva of Bothriomyrmex enigmaticus sp. nov., profile view.

7

European Journal of Taxonomy 211: 1-12 (2016)

Petiolar node in profile scale-like and strongly inclined anteriorly, with the anterior face much shorter

than the posterior face (Fig. 2c). Ventral margin of petiole with a large lobe. Several setae present on the

posteroventral margin of lobe.

Second, third and fourth tergites of gaster with long, erect setae arising from the middle of the tergite:

two on the second, six on the third and fourth. First four sternites of gaster also bearing two long setae

similar to those found on the tergites. Pubescence similar to the rest of the body, but becoming longer on

the posterior margins of the sclerites.

Uniformly light brown; coxae and legs somewhat lighter.

Larva description

Shape dolichoderoid. Body with two ventrolateral protuberances on prothorax which are fused ventrally

by a narrow ridge; setae very short and limited to the prothorax; 8 pairs of spiracles. (Fig. 4).

Key to Neotropical Bothriomyrmex species based on workers

1. Palp formula 4,3 (see fig. 3d in Dubovikoff & Longino 2004); medial lobe of clypeus strongly

projecting beyond the lateral lobes; anterior margin evenly rounded to flat, never concave medially;

paired long setae on anterior margin of medial lobe separated by less than 2 /3 of their length (Fig. 2a)

. B. paradoxus (Dubovikoff & Longino, 2004)

- Palp formula 2,3 (Fig. 2d); medial lobe of clypeus weakly projecting; anterior margin flat to broadly

concave medially; paired long setae on anterior margin of medial lobe separated by 2 V of their

length or more (Fig. 2b) . B. enigmaticus sp. nov.

Fig. 5. Known distribution of Bothriomyrmex paradoxus (Dubovikoff & Longino, 2004) and

B. enigmaticus sp. nov.

8

PREBUS M. & LUBERTAZZID., Caribbean Bothriomyrmex

Distribution and ecology

Bothriomyrmex enigmaticus sp. nov. is known from one collection on the north side of the island of

Hispanola (Fig. 5). The sampling took place in July 2015 during a Museum of Comparative Zoology

expedition to the Dominican Republic. The new ant species was discovered while based at a guardhouse

in the Toro Palomo sector on the SE side of the Loma Guaconejo Scientific Reserve, near the village

of La Peonia, at 195 m in elevation. M. Prebus collected a partial nest containing workers and brood

from decomposing wood at the top of a 1.5 m tall live sapling in the middle of a path leading west from

the guardhouse. The habitat was scrubby secondary growth, roughly 100 meters from a more mature

secondary lowland moist forest. Tapinoma litorale (Wheeler, 1905) was abundant in this habitat; due

to the superficial similarity of these two species, Bothriomyrmex enigmaticus sp. nov. was mistaken for

T. litorale in the field. The nest was not mixed; it consisted solely of B. enigmatus.

Taxonomic notes

Dubovikoff & Longino (2004) came to the conclusion that Bothriomyrmex paradoxus is a member of

the Palearctic Bothriomyrmex s.s. based on palp formula and wing venation. They also noted that it

doesn’t fit perfectly into species groups in this region: a deeply impressed metanotal groove is found

in the B. gib bus (Soudek, 1925) group, but the gyne of B. paradoxus also has short, suberect setae on

the mesosoma and gaster, which is typical of the B. syrius (Forel, 1910) group. With its large eyes,

deep metanotal groove, and nesting preference, B. enigmaticus sp. nov. appears to be closely related

to B. paradoxus. However, it diverges from the latter in several characters, most notably palp formula.

Referring to Shattuck (1992), it appears that a palp formula of 6,4 is probably pleisiomorphic in the

Dolichoderinae. Wit hin the tribe Bothriomyrmecini there is a trend in palp reduction in Arnoldius ,

Bothriomyrmex and Chronoxenus (Table 1) that becomes apparent when they are compared with

Loweriella (Shattuck, 1992) and Ravavy (Fisher, 2009), which have palp formula 6,4 and 6,3. It is

possible that a palp formula of 4,3 is plesiomorphic in the clade of ( Bothriomyrmex + Chronoxenus +

Arnoldius ), with the latter two genera having undergone subsequent reductions. Within other genera

of the Dolichoderinae, e.g., Technomyrmex (Mayr, 1872) and Azteca (Forel, 1878), palp formula can

be highly variable, making generic diagnoses based on this character alone problematic. While the

reproductives (and therefore the wing venation) of B. enigmaticus sp. nov. remain unknown, if the

worker were to be classified based on the current generic diagnoses, one might be inclined to place it in

Chronoxenus based solely on palp formula. However, the similarities between B. enigmaticus sp. nov.

and B. paradoxus mentioned above are striking, suggesting that these two species are sister taxa and

palp formula is a labile character.

Placement in Bothriomyrmecini and biogeographic notes

Material examined

Bothriomyrmex paradoxus (Dubovikoff & Longino, 2004)

Paratype worker

COSTA RICA: Heredia, La Virgen, 10 km SE La Virgen, 10.33333° N, 84.08333° W, 500 m, 16 Feb.

2003, wet forest treefall gap, under epiphytes, J. Longino #JTL4938 [UCDC: JTLC000003511]

Chronoxenus wroughtoni (Forel, 1895)

CHINA: Hong Kong, 1 worker, N.T., Kadoorie Institute Shek Kong Centre, 22.42833° N, 114.11455° E

± 7 m, 220 m, 3 Jul. 2011, secondary rainforest, under stone, RS. Ward #PSW16602 [UCDC:

CASENT0220768].

9

Table 2. Worker morphometric data adapted from Seifert (2012), comparing Bothriomyrmexparadoxus (Dubovikoff & Longino, 2004), B. enigmaticus

sp. nov., Chronoxenus wroughtoni (Forel, 1895) and Arnoldius (Dubovikoff, 2005) AU01 to Bothriomyrmex (Emery, 1869) species from the western

Palearctic. Abbreviations: n = number of nest samples; i = number of individuals. Data is expressed as arithmetic mean ± standard deviation [lower

extreme, upper extreme].

European Journal of Taxonomy 211: 1-12 (2016)

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10

PREBUS M. & LUBERTAZZID., Caribbean Bothriomyrmex

Arnoldius (Dubovikoff, 2005) AU01

AUSTRALIA: 1 worker, Queensland, Smithfield Conservation Park 16.8167° S, 145.6833° E,

70 m, 9 Mar. 2006, rainforest, sifted litter (leaf mold, rotten wood), A. Lucky #ALC339-1 [UCDC:

CASENT0106155]; 1 worker, New South Wales, Kanangra, Boyd National Park, 33°59' S, 150°07' E,

1050 m, 19 Jul. 1975, dry sclerophyll forest, under stone, PS. Ward #PSW1000 [UCDC]; 1 worker,

Queensland, Smithfield Conservation Park, 16°49' S, 145°41' E, 70 m, 17 Aug. 2006, rainforest,

ex. rotten log, PS. Ward #PSW15681 [UCDC],

In Table 2 we compare B. enigmaticus sp. nov., B. paradoxus , Chronoxenus wroughtoni and an undes¬

cribed species of Arnoldius to data collected from a morphometric study of Palearctic Bothriomyrmex

conducted by Seifert (2012). In this context, some very general observations can be made about the

morphological affinities of B. enigmaticus sp. nov.: the eyes of B. enigmaticus sp. nov., relative to

cephalic size are the largest in this dataset, similar to the large-eyed B. paradoxus , C. wroughtoni , and

B. corsicus. The antennal scapes are short as well, a feature shared by B. paradoxus , C. wroughtoni and

A. AU01. Finally, the propodeum is compact and delineated by a deep metanotal groove, much like

B. paradoxus , C. wroughtoni , A. AU01, and to a lesser degree B. corsicsus.

Discussion

This is the first record of Bothriomyrmex from the Caribbean region, and the second species to be

described from the Americas. Until very recently, this genus was considered to be limited strictly

to the Palearctic region. An estimation of the phylogeny, biogeography and divergence times of the

Dolichoderinae (Ward et al. 2010) suggested that the split between B. paradoxus and the Palearctic

species, B. sounder si (Santschi, 1922), occurred between 5 and 15 my a, excluding a simple explanation

of vicariance by continental drift. One hypothesis that could reconcile the current distribution and

phylogeny of this group is that characters such as short antennal scapes, compact propodeum and deep

metanotal groove are plesiomorphic to ( Bothriomyrmex + Chronoxenus + Arnoldius ), while having

large eyes is a trait that is plesiomorphic to ( Chronoxenus + Bothriomyrmex ), which have subsequently

become reduced in some species. If this is the case, the most plausible explanation for the current

distribution is that Bothriomyrmex was once much more broadly distributed in the Americas, and went

extinct in the Nearctic part of its range in recent t im es.

Acknowledgements

We wish to thank Grupo Jaragua for their assistance: Ernst Rupp for logistical support. Hector Andujar

for managing our travels and his field assistance, Jose Luis Castillo for his translation skills and his

field assistance, and Jairo Isaac Matos for his field assistance. Santo Navarro, Universidad Autonoma

de Santo Domingo, and Gabriel de los Santos, Museo Nacional de Historia Natural, helped in numerous

ways to insure our expedition and research went smoothly. Rafael Emilio Gomez, provincial director

of the Ministerio de Medio Ambiente y Recursos Naturales, graciously provided us with lodging and

the assistance of his staff during our visit to Loma Guaconejo Scientific Reserve. We thank them and

the Santo Domingo office of the Ministerio de Medio Ambiente y Recursos Naturales for providing

the permits and permissions required for this research. David Lubertazzi wishes to thank Stefan Cover

(Museum of Comparative Zoology) and Matthew Prebus wishes to thank Phil Ward (UC Davis) for their

support with this study and with all of our myrmecological endeavors. The UC Davis Department of

Entomology and Nematology provided material support to Matthew Prebus that helped make this work

possible. The Green Fund and the Museum of Comparative Zoology provided funding and other support

for the field expedition.

11

European Journal of Taxonomy 211: 1-12 (2016)

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Manuscript received: 13 November 2015

Manuscript accepted: 9 March 2016

Published on: 11 July 2016

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands.

12