European Journal of Taxonomy 85: 1-10

http://dx.doi.org/10.5852/ejt.2014.85

ISSN 2118-9773

www. europeanj oumaloftaxonomv. eu

2014 • Fernandez F., Feitosa R.M. & Lattice J.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

urn:lsid:zoobank.org:pub:22BEA612-F21A-4FA6-853F-ED9C430649C3

Kempfidris, a new genus of myrmicine ants from the Neotropical region

(Hymenoptera: Formicidae)

Fernando FERNANDEZRodrigo M. FEITOSA 2 & John LATTKE 3

1 Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogota D.C.,

Colombia. E-mail: [email protected]

um:lsid:zoobank.org:author:6F96DA0B-5889-40F4-A946-DBB05FB77F7B

2 Departamento de Zoologia, Universidade Federal do Parana, Caixa Postal 19020, 81531-980,

Curitiba, PR, Brazil. E-mail: rsmfeitosa@gmail com

urn:lsid:zoobank.org:author:443A72A7-196F-47FF-A02D-0B5F4B48D8B3

3 Universidad Nacional de Loja, Direccion de Investigacion — BioS, Ciudad Universitaria,

La Argelia, Loja, Ecuador. E-mail: piquihuye@, email,com

urn:lsid:zoobank.org:author:7AE0D94C-619B-4E77-B16F-3F18731EE390

Abstract. The new genus Kempfidris gen. nov. is described based on the workers of a single species,

K. inusualis comb, nov., from Brazil, Ecuador, and Venezuela. Kempfidris inusualis comb. nov. was

originally described by Fernandez (2007) and provisionally placed in Monomorium awaiting a better

understanding of the internal relationships in Myrmicinae. Kempfidris gen. nov. has a series of distinctive

morphological characters including the mandibular configuration, vestibulate propodeal spiracle,

propodeal carinae, and cylindrical micro-pegs on the posteromedian portion of abdominal tergum VI

and anteromedian portion of abdominal tergum VII. This last trait appears to be autapomorphic for the

genus.

Key words. Myrmicinae, new genus. Neotropical region, taxonomy

Fernandez F., Feitosa R.M. & Lattice J. 2014. Kempfidris, a new genus of myrmecine ants from the Neotropical

region (Hymenoptera: Formicidae). European Journal'of Taxonomy 85:1-10. http://dx.doi.org/10.5852/eit.2014.85

Introduction

Due to its large diversity, Myrmicinae Lepeletier, 1835 is the most challenging subfamily in myrmecology

regarding its internal relationships and the search for a stable tribal arrangement. Much of the current

classification follows the initial proposals of Emery (1921) and Wheeler (1922), with important

modifications by Bolton (2003). However, many tribes and some genera are obviously artificial, and

given the considerable amount of ongoing research, both basic and applied, involving many taxa from

this group, it is important to begin to clarify the internal relationships of this subfamily.

The solenopsidine tribe group, proposed by Bolton (2003) for Solenopsidini Forel, 1893 and Stenammini

Ashmead, 1905, is partly diagnosed by a clypeus with a posterior constriction moderately to greatly

inserted between the frontal lobes and an elevated and narrow median area, which is usually bicarinate.

The same author recognized the Solenopsidini, including the Solenopsis group, characterized by the

European Journal of Taxonomy 85: 1-10 (2014)

presence of a seta on the anteromedian clypeal margin, and the Carebara group, which lacks the clypeal

seta. Subsequent modifications to this proposal include the creation of the tribe Adelomyrmecini

Fernandez, 2004 (, Adelomyrmex Emery, 1897, Baracidris Bolton, 1981 and Cryptomyrmex Fernandez,

2004) and an expansion of the concept of Monomorium Mayr, 1855 in the Neotropics (Fernandez

2004, 2007). However, there have been no phylogenetic studies evaluating either these proposals or the

monophyly and relationships of the solenopsidine group.

Some central problems remain in defining Monomorium and the relationships between the satellite taxa

proposed throughout its long taxonomic history, including Monomorium inusuale Fernandez, 2007.

Recent molecular and morphological phylogenetic studies of ants as a whole have been focused at the

subfamily or tribal level, beyond the scope of issues at the generic level (Moreau et al. 2006; Brady et al.

2006; Keller 2011; Moreau & Bell 2013). Solenopsidini does appear to be polyphyletic in Moreau et al.

(2006), but the status of Monomorium remains unclear. According to Brady et al. (2006), Solenopsidini

could be monophyletic, but with the inclusion of Myrmicaria Saunders, 1842, whereas the status of

Monomorium remains unknown. In Moreau & Bell (2013), Solenopsidini appears to be monophyletic if

Myrmicaria is included, and Monomorium is apparently polyphyletic. Even though these studies did not

address the generic level, they do make an unequivocal case for the poor resolution and instability of the

relationships between the tribes and genera proposed within the Myrmicinae.

Currently, the Ant Tree of Life project (AToL) team is engaged in the final stages of preparing a

presentation of the internal phylogeny of Myrmicinae, and preliminary results corroborate the artificial

nature of the cosmopolitan and morphologically diverse genus Monomorium , already suspected by

various authors (Bolton 1987; Heterick 2003). In 2007, Fernandez described two new species for the

Neotropics, including the morphologically peculiar Monomorium inusuale, based on workers collected

in forested areas of Brazil (Amazonas and Bahia) and Ecuador. Workers of M. inusuale present a very

unusual combination of morphological characters, which led Fernandez to question the position of the

species within the genus. However, M. inusuale was tentatively assigned to Monomorium until the limits

of the heterogeneous myrmicine tribes could be clearly defined.

A reconsideration of the characters defining M. inusuale, and its position amongst the solenopsidines, has

strengthened the case for recognizing this taxon as a distinct lineage in this group. Here we describe this

new monotypic myrmicine genus and provide updated information on its distribution in the Neotropical

region. We consider this as one of the many necessary steps to seeking a more stable classification in

Myrmicinae.

Material and methods

The materials studied during this investigation are deposited in the following institutions:

CASC = California Academy of Sciences, San Francisco, CA, USA

DZUP = Cole£ao Entomologica Pe. Jesus Santiago Moure, Universidade Federal do Parana,

Curitiba, PR, Brazil

ICN = Instituto de Ciencias Naturales, Universidad Nacional de Colombia, BogotaD.C., Colombia

MZSP = Museu de Zoologia, Universidade de Sao Paulo, SP, Brazil

Scanning electron microscope (SEM) photographs (Fig. 1) were taken at the University of Alberta in

Edmonton by Danny Shipley and George Ball. High resolution images (Fig. 2) were obtained using a

Leica M125 stereomicroscope attached to a Leica DFC 295 video camera. Photos were combined using

Zerene Stacker software at the DZUP. Images were then processed as TIFF files in Adobe® Photoshop

CS5® to enhance parameters of brightness and contrast. Pictures of the Venezuelan specimen (Fig. 3)

were taken from AntWeb ( www.antweb.org ) .

FERNANDEZ F., FEITOSAR.M. & LATTKE J., New ant genus from the Neotropical region

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Suborder Apocrita Latreille, 1810

Superfamily Vespoidea Latreille, 1802

Family Formicidae Latreille, 1809

Subfamily Myrmicinae Lepeletier, 1835

Tribe incertae sedis

Kempfidris gen. nov.

Figs 1-3

urn:lsid:zoobank.org:act:B9B9FBA4-E0A5-44A2-BDDl-3C9ADA644AD9

Type species

Monomorium inusuale Fernandez, 2007

Etymology

It is with great pleasure that we dedicate this new genus to Father Walter W. Kempf (1920-1976) (Fig.

4) for his brilliant, but somewhat short career in ant taxonomy, with special reference to the Neotropical

fauna. Besides the many monographs and descriptions of new taxa, Kempf provided the first catalog of

Neotropical ants (Kempf 1972), an obligate reference (until recently) for generations of myrmecologists.

Description (worker)

Antenna 12-segmented, with a well-defined 3-segmented club. Eye reduced to almost indistinct

ommatidia, placed just anteriorad of head midlength. Frontal carinae and antennal scrobes absent.

Mandible with four teeth: three apical teeth separated from isolated basal tooth by large diastema. Palpal

formula 2,1. Clypeus projected anteromedially to form truncate lobe, lobe laterally bluntly angular and

medially weakly concave, with distinct median seta and several surrounding setae. Median lobe of

clypeus strongly bicarinate, carinae defining elevated area that converges posteriorly to form narrow

strip between frontal lobes. Pronotum with anterolateral crest that extends dorsally, delimiting narrow

anterior portion of pronotum. Metanotal groove very deep, well-marked. Bulla of metapleural gland

large. Propodeum unarmed, declivity surrounded dorsolaterally by a crest, higher laterally than dorsally

and joining the small metapleural lobe. Posteromedian portion of abdominal tergum VI and anteromedian

portion of abdominal tergum VII with several minute, cylindrical micro-pegs, each bearing a hair on

apex, though apparently broken in some.

Remarks

Kempfidris inusualis comb. nov. was provisionally described in Monomorium (Fernandez 2007), but with

many doubts due to the poor state of knowledge of the internal relationships in Myrmicinae. Fernandez

commented on a suggestion by Barry Bolton (pers. comm.) that K. inusualis comb. nov. might be a

member of Stena mmini due to its overall resemblance with members of that tribe. Traits of the frontal

lobes and toruli, as well as the clypeal posterior border, would place K. inusualis comb. nov. in the

Stenammini as diagnosed in Bolton (2003: 58). However, the first gastral tergite clearly overlaps the first

sternite on its ventral surface, and the presence of a single medial clypeal seta indicates that the taxon

belongs in the Solenopsidini, again sensu Bolton (2003: 59). We presently prefer to leave Kempfidris

gen. nov. as an incertae sedis genus within the Myrmicinae until Ward and collaborators publish the

conclusions, including a tribal arrangement, of their ongoing research (PS. Ward, pers. comm.).

European Journal of Taxonomy 85 : 1-10 ( 2014 )

A reconsideration of this species, given the series of unusual characters it has compared to other

myrmicines, argues for recognising it as a distinct lineage. Leaving it in Monomorium only increases

the difficulties of defining a monophyletic Monomorium , thus confounding the present situation within

those groups. The strongly reduced eyes and the long diastema between the three apical teeth and

the basal tooth do not appear to be co mm on in myrmicines. Kempfidris gen. nov. is reminiscent of

Cryptomyrmex regarding the mandibular configuration (four teeth with a diastema), clypeal structure

(elevated median area), single median seta along the anterior clypeal margin, and reduced number (2)

of maxillary palp segments. However, Cryptomyrmex has a different antennal structure (12 segments,

the last two forming a club), modified hairs along the mandibular masticatory margin and subcutaneous

ommatidia. Reduced eyes can be found in the African fossulatum- group of Monomorium (Bolton 1987),

as well as in Carebarella.

Fig. 1. Kempfidris inusualis (Fernandez, 2007) comb, nov., worker from Benjamin Constant, Amazon,

Brazil (ICN). SEM photographs. A. Head in full face view. B. Head in dorsal oblique view. C. Mesosoma

in lateral view. D. Apex of metasoma showing the cylindrical micro-pegs.

FERNANDEZ F., FEITOSAR.M. & LATTKE J., New ant genus from the Neotropical region

The most outstanding feature of Kempfidris gen. nov. is the series of minute, hair-bearing tubercles

or cylindrical pegs on the abdominal apex. Most are concentrated on the anteromedian portion of the

pygidium and some on the posteromedian portion of abdominal tergite VI, a position that would coincide

with the position of the pygidial gland, which opens between abdominal tergites VI and VII (Billen

2009). The structure of the micropegs with their associated hairs also hints at some sort of glandular

function, or possibly a mechano-reception function during stinging, but a more convincing explanation

will only be possible after a histological study. Whatever the function of these tubercles, they appear

to be an autapomorphic structure, absent in other Myrmicinae and probably in other ants as well. In

Fig. 2. Kempfidris inusualis (Fernandez, 2007) comb, nov., worker from Porto Velho, Rondonia, Brazil

(DZUP). High-resolution images. A. Head in full face view. B. Body in dorsal view. C. Body in lateral

view.

European Journal of Taxonomy 85: 1-10 (2014)

ants, the most structurally similar cuticular projections can be found throughout most of the body of

some species in the formicine genus Echinopla Smith, 1857, except on the pygidium, and additionally

on the gastral apex of an undescribed species of Strumigenys Smith, 1860 (Myrmicinae: Dacetini).

Given that these aforementioned taxa are not closely related to Kempfidris gen. nov., their structures

are probably not homologous, but perhaps convergent evolution could be considered, especially in the

case of the dacetine ant. Other specialized pygidial structures found in ants are the denticles or spines

of Cerapachyinae and the large, upward-curving teeth in Pachycondyla crassinoda (Latreille, 1802)

workers, but their position and form are very different. Most members of the solenopsidine group are

smooth, with little sculpturing, but this species presents a moderate amount of sculpturing on the head,

mesosoma, petiole and postpetiole.

Fig. 3. Kempfidris inusualis (Fernandez, 2007) comb, nov., worker from Maroa, Amazonas, Venezuela,

specimen CASENT0217050. High-resolution images from AntWeb. A. Head in full face view. B. Body

in dorsal view. C. Body in lateral view. Images by Erin Prado.

FERNANDEZ F., FEITOSAR.M. & LATTKE J., New ant genus from the Neotropical region

Kempfidris inusualis (Fernandez, 2007) comb. nov.

Figs 1-3, 5

Monomorium inusuale Fernandez, 2007: 135, figs 1-2 (worker).

Material examined

Holotype

Worker, Brazil, Bahia, Barrolandia, 16-23 Jul. 1994, S. Lacau (CEPLAC = Laboratorio de Formigas,

Centro de Pesquisas de Cacau, Ilheus, Bahia, Brazil).

Paratypes

1 worker, same data as holotype (ICN); 3 workers, Brazil, Amazonas, Benjamin Constant, 21 Nov.

1962, W.L. Brown Jr. (MZSP); 2 workers, Ecuador, Napo, Limoncocha, 1973, M. Rettenmeyer (MZSP,

BMNH). For measurements, see Fernandez (2007).

Additional material

BRAZIL: 12 workers, Rondonia, Porto Velho, Mutum, M7P1, 9°35’44” S 65°04’00” W, 8-21 Sep.

2010, R.R. da Silva & R.M. Feitosa (MZSP); 3 workers, same data (DZUP).

VENEZUELA: 2 workers, Amazonas, Maroa, 120 m, 2°43 , 48”N 67°32 , 34” W, 4 Sep. 2006, J.E. Lattke,

JEL3031 (CASC, 1;ICN, 1).

Fig. 4. Father Walter Kempf (1920-

1976). Photographer unknown;

files of the Franciscan Order in Sao

Paulo, Brazil. Available from the

Global Ant Project website: http://

gap. entclub. org/taxonomists/Kempf/

index.html

European Journal of Taxonomy 85: 1-10 (2014)

Description (worker)

Head longer than wide. Sides of head subparallel, slightly wider anteriorly, broadly rounded into

posterior border, which is more or less straight. Antennal insertions relatively close together, separated

by no more than three times the apical scape width. Scape fails to reach vertexal border.

In lateral view, promesonotum slightly convex, mesonotal margin weakly convex to almost straight.

Metanotal groove very deep, well-marked. Propodeal spiracle circular, opened posteriorly. Petiole with

peduncle and node well differentiated, the node with subparallel anterior and posterior faces, dorsum

convex. Petiolar spiracle at base of node. Petiolar ventral process shaped as rounded, anteroventrally

directed tooth. Postpetiole subcampaniform, with a conspicuous transverse, ventral carina. Mandibles,

most of promesonotum, dorsum of petiole and postpetiole, and gaster smooth and shining. Head with

longitudinal, irregular rugulae mixed with dense foveae. Posterior portion of promesonotum with feeble,

short rugulae. Most of mesopleura with irregular, short, longitudinal striae, mixed with fine reticulation.

Fig. 5. Distribution map for Kempfidris inusualis (Fernandez, 2007) comb. nov.

FERNANDEZ F., FEITOSAR.M. & LATTKE J., New ant genus from the Neotropical region

Most of propodeum and sides of petiole and postpetiole densely reticulated. Declivity of propodeum

with several fine transverse carinae, the most posterior more marked. Moderately erect pilosity on head,

promesonotum, petiole, postpetiole and gaster, little on propodeum. Scapes with several erect hairs.

Longer hairs about 0.15 mm, those of head shorter. Body light brown to black, appendages lighter.

Queen and male: Unknown.

Distribution

Brazil, Ecuador and Venezuela (Fig. 5).

Discussion

Despite the fact that this species is broadly distributed in South America (Fig. 5), records of its occurrence

are extremely scarce. Until recently, Kempfidris inusualis comb. nov. was known only from the type

series examined by Fernandez (2007) and a nest series collected by John Lattice in 2006. Unfortunately,

the bulk of this material was left in Venezuela upon Lattice’s recent move from the country. However,

in 2010 a field expedition (which included R.M. Feitosa) to the southern limits of the Brazilian Am azon

Forest (state of Rondonia, near the border with Bolivia) collected a series of K. inusualis comb. nov.

workers extracted from a Winkler sample.

Despite its broad distribution, specimens of K. inusualis comb. nov. are relatively uniform in size and

general aspect. As already mentioned by Fernandez (2007), the main differences regarding geographic

variation involve slightly distinct propodeal shapes, such as a shorter dorsal face in the Ecuadorian

specimens. Specimens from Rondonia, Brazil, differ from other conspecific workers by their darker

color (almost black) and more prominent propodeal crests that form small denticles. Given the similarity

among the samples examined, we decided to consider these morphological differences as intraspecific

variation.

Almost nothing is known about the natural history of K. inusualis comb. nov. Most of the specimens

studied were apparently obtained from leaf-litter samples. The nest series collected by J. Lattice in the

Venezuelan Amazon was obtained from a rotten stick, also harboring a termite nest, on the ground next

to an airstrip. The habitat there is open scrub to low trees on white sandy soil that is seasonally flooded by

dark, colored waters. Interestingly, all the known samples were collected between July and September,

suggesting a more intense activity in this period, which coincides with the low level of the rivers in the

Amazon Basin. All things considered, this could also be a collecting artifact, as the aforementioned

period approximately coincides with academic holidays in many universities and represents a lower

probability of conflict between lectures and field trips.

Acknowledgements

Dr. Philip Ward (University of California at Davis) for generously sharing unpublished information on

the internal phylogeny of Myrmicinae. Bru nn o Bueno, Bruno Ferrari, and Gabriel Melo (DZUP) for

kindly helping with the preparation of the automontage images. The Secretary of Higher Education,

Science, Technology and Innovation (SENESCYT) of the Government of Ecuador for supporting

J. Lattice s research. To the topic editor of the European Journal of Taxonomy and two anonymous

referees for their appropriate corrections and suggestions; one of them provided useful comparative

notes regarding the micro-pegs on the abdominal apex.

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Manuscript received: 14 March 2014

Manuscript accepted: 9 May 2014

Published on: 26 May 2014

Topic editor: Koen Martens

Desk editor: Danny Eibye-Jacobsen

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; National History Museum, London, United

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

Denmark, Copenhagen, Denmark.