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NOTES ON GEOGRAPHIC DISTRIBUTION Check List 14 (2): 291-296

First records of the minor pest termite Eucryptotermes hagenii
(Miiller, 1873) (Blattodea, Termitoidae, Kalotermitidae) from the
Chaco Dominion in Argentina

Maria C. Godoy, Juan M. Coronel, Giovana M. Annoni, Clara Etcheverry, Enrique R. Laffont

Universidad Nacional del Nordeste, Facultad de Ciencias Exactas y Naturales y Agrimensura, Biologia de los Invertebrados, Av. Libertad 5470,
Corrientes, Argentina.
Corresponding author: Maria C. Godoy, mcgodoy@exa.unne.edu.ar

Abstract

Eucryptotermes Holmgren, 1911, comprises 2 species of Neotropical drywood termites with phragmotic head soldiers.
We report the presence of Eucryptotermes hagenii (Miller, 1873) in native forests of the Chaco Dominion and Chaco
Province, enlarging its distribution and recording the genus and species for the first time in Argentina. Eucryptotermes
hagenii abundance was estimated as intermediate by standardized sampling. Eighteen morphometric characters were
measured in soldiers and alates, and 9 of them provided new data. The colonies, located inside living trees trunks and
dead wood, were composed of reproductives, soldiers, pseudergates, and immatures.

Key words
Insecta; drywood termites; South America; distribution.

Academic editor: Pedro Guilherme Barrios de Souza Dias | Received 5 September 2017 | Accepted 12 December 2017 | Published 2 March 2018

Citation: Godoy MC, Coronel JM, Annoni GM, Etcheverry C, Laffont ER (2017) First records of the minor pest termite Eucryptotermes
hagenii (Miller, 1873) (Blattodea, Termitoidae, Kalotermitidae) from the Chaco Dominion in Argentina. Check List 14 (2): 291-296. https://doi.
org/10.15560/14.2.291

Introduction pest status. Most of records of E. hagenii are from natu-
ral habitats in southeastern Brazil, but it is also known
from urban areas, such as Curitiba and Blumenau, which
wood termites” that live in trees and pieces of wood. indicates that it could be an occasional structural pest

Soldiers have truncated heads with a remarkable degree (Constantino 1997, 1998). In the city of Sao Paulo,
of phragmosis (Krishna 1961). This genus has only 2 where the most significant termite damage is caused by
described species, the type species, E. hagenii (Muller, Cryptotermes Banks, 1906 and Coptotermes Wasmann,
1873), and E. breviceps Constantino, 1997. These two _ 1896, E. hagenii is considered to be a minor urban pest
species were previously known only in Brazil, inthe Bra- (Fontes 1995, Constantino 2002). From a cytogenetic
zilian and Chaco Subregions, respectively (Araujo 1977, and evolutionary point of view, this species is unusual
Constantino 1998, Krishna et al. 2013). because it has the lowest number of chromosomes so far

The published data on E. hagenii are few and include reported for termites, with 2n = 22 (Martins and Mesa
some biological and cytogenetic aspects, as well as its 1995).

Eucryptotermes Holmgren, 1911, is a little-known and
highly specialized endemic Neotropical genus of “dry-

Copyright Godoy 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.

292

Check List 14 (2)

0 320640 960 km

Figure 1. Geographical distribution of the genus Eucryptotermes (South America). Circle: E. breviceps; squares: E. hagenii.

We report for the first time the presence of E. hagenii
in natural habitats in the Chaco Dominion in Argentina.
Our new records enlarge the known range of this spe-
cies and also provide new information on its biology and
morphometry.

Methods

The samples were collected as part of a research project
on termite communities in the Humedales Chaco Ramsar
Site at the Los Chaguares Reserve (Colonia Benitez,
Chaco province, Argentina) between September 2016
and February 2017 (Fig. 1) (27°20.00'S, 058°58.03'W).
The Reserve (19 ha) is located in an area with humid
mesothermal climate and little winter rainfall, where
the average annual temperature is above 18 °C. Biogeo-
graphically, it belongs to the Chaco Subregion, Chaco
Dominion, Chaco Province, Eastern Chaco District or
Humid Chaco Ecoregion and is characterized by the
presence of grasslands, floating islands (“embalsados’’),

gallery forests, and xerophytic forests (Morrone 2014).
The vegetation in the study area is a riverbank forest with
trees up to 15 m high (Cabrera and Willink 1973, Cabrera
1976).

The standardized sampling protocol for termites
(Jones and Eggleton 2000) was applied, consisting of 2
transects, each 100 x 2 m, that are divided into 20 contig-
uous sections (5 x 2 m). Termites were collected in each
section (1 person/h) in various microhabitats, including
fallen tree trunks and branches, mounds, and arboreal
nests. Twelve samples of surface soil (12 x 12 cm, 10 cm
deep) were excavated in each section. The total number
of termite encounters per transect was counted.

The abundance of E. hagenii was estimated in the
surveyed areas, defining their occurrence, dominance
and abundance patterns according to the categories estab-
lished by Florencio and Diehl (2006). To determine the
occurrence pattern, the species are categorized as rare
(R: present in 1-10% of sections), sporadic (S: 11-40%
of sections), common (C: 41—70% of sections), frequent

Godoy et al. | Eucryptotermes hagenii in Chaco, Argentina

(F: 71-99% of sections), and constant (CS: 100% of sec-
tions). The following formula was applied: OP = numbers
of sections where the species was found = 100/total num-
ber of evaluated sections. For the dominance pattern, the
species are classified as rare (R: 1-10% of encounters),
accessory (A: 11-49% of encounters), and dominant (D:
50—100% of encounters), according to the formula DP =
number of encounters of each species < 100/total number
of encounters. The combination of both patterns 1s used
as an indicator of the abundance of each species, accord-
ing to the following categories: common (C: species with
CS and D patterns), intermediate (I: species with R, S,
C or F and A and species with S, C, F or CS and R),
and rare (R: species with R in occurrence and dominance
patterns).

Specimens were identified by M.C. Godoy (first
author) using taxonomic keys and descriptions (Muller
1873, Silvestri 1903, Holmgren 1911, Krishna 1961,
Constantino 1997, 1999) and by comparisons with sam-
ples of E. hagenii from Sao Paulo, Brazil, which were
determined by L.R. Fontes (Termite collection, Facultad
de Ciencias Exactas y Naturales y Agrimensura, Uni-
versidad Nacional del Nordeste, Corrientes, Argentina;
FACENAC).

Voucher specimens were preserved in 80% ethanol
and deposited in the FACENAC (registration numbers
2512-2518). The specimens were collected under the
authorization of the Direccién de Fauna, Parques y
Ecologia, Chaco Province.

In soldiers and alates, 18 morphometric characters
(Roonwal 1970, Scheffrahn and Krecek 1999) were mea-
sured with a micrometric eyepiece attached to a Leica
EZ4 stereomicroscope. Microphotographs were taken
with a Canon EOS Rebel T3 digital camera coupled to
the microscope. For scanning electron microscopy, the
specimens were dehydrated in a conventional ethanol
series, air-dried, gold-coated, and photographed with the
SEM Jeol 5800 LV from the UNNE Microscopy Service.
The map was elaborated by using QGIS Essen version
2.14 (QGIS 2016).

Results

New records. Argentina, Chaco, Colonia Benitez (27°
20.00'S, 058°58.03'W): MCG and ERL, 24 November
2016, FACENAC 2512, 7 alates, several soldiers and
pseudergates; GMA, 23 November 2016, FACENAC
2513, 1 soldier and several pseudergates,; MCG and
ERL, 24 November 2016, FACENAC 2514, 1 soldier
and several pseudergates; JMC and ERL, 24 November
2016, FACENAC 2516, several pseudergates; JMC and
CE, 24 November 2016, FACENAC 2517, 1 dealate, 4
soldiers and several pseudergates; MCG, 24 November
2016, FACENAC 2518, 2 dealates, 3 soldiers and sev-
eral pseudergates; JMC and GMA, 23 November 2016,
FACENAC 3155, several prealates and pseudergates;
MCG and ERL, 23 november 2016, FACENAC 3157, 1
soldier and several prealates.

293

Identification. Soldiers of £. hagenii (Figs 2-6) have a
dark brown, short, and broad sub-truncated or phragmotic
head, with almost straight parallel sides and a slight con-
striction at the antennae level. The frontal region of the
cephalic capsule is extended forward, forming a sharp
circular ridge or frontal flange located dorsally between
the vertex and the frons and between the frons and the
genae on both sides. The antennae are inserted at about
one-third or one-half of the length of the head from the
front sides (Fig. 4). The mandibles are partially covered
by the cephalic capsule in dorsal view, and its inner edge
is slightly serrated without notable teeth. The pronotum is
brownish yellow with its anterior margin rising abruptly
in profile, emarginated, and strikingly serrated (Fig.
5). The alates of E. hagenii have a uniformly brownish
yellow coloration and transparent wings (Fig. 7). The
pronotum is narrower than the head. The anterior margin
of the forewing scale is almost straight or very slightly
convex and forewings have all major veins emerging
independently at the wing suture. The 18 morphometric
characters measured on E. hagenii soldiers and alates are
presented in Table 1.

Eucryptotermes breviceps differs from E. hagenii by
its smaller soldiers and imagoes and by having only 2
tibial spurs on the fore-tibiae. Soldiers of E. breviceps
also have a shorter and strongly phragmotic head, with a
much larger frontal extension covering most of the man-
dibles in dorsal view (Constantino 1997).

Other kalotermitid genera detected in Argentina, with
soldiers showing varying degrees of cephalic phragmo-
sis are Cryptotermes, Glyptotermes Froggatt, 1897, and
Tauritermes Krishna, 1961, but the development of their
frontal ridge is smaller than in Eucryptotermes (Silves-
tri 1903, Torales et al. 1997, Roisin 2003). In addition,
unlike them, the antennae of Eucryptotermes are inserted
at about one-third or one-half of the head length from the
front sides.

Discussion

In our surveys of termites in native forests of the Hume-
dales Chaco Ramsar Site, E. hagenii was collected and
identified from the Chaco Dominion, Chaco Province,
Chaco Humid District for the first time (Fig. 1). In
this biome, £. hagenii 1s a sporadic species, because 6
colonies were detected in the 2 transects (12.5% of the
analyzed sections). With regard to dominance, it was an
accessory species (15.79%), so that the abundance of F.
hagenii in the analyzed forests was categorized as inter-
mediate. Other 2 colonies of the species were detected in
the Reserve during complementary samplings.

We found only 2 nests in Colonia Benitez that had 1 or
2 dealate reproductives. Presumably these are members
of the royal couple and suggest that the colonies were
monogynous. In November, 7 mature alates were found
in a single nest. The mature alates found in November
indicate that swarming might take place in late spring or
early summer. All colonies were also composed of sol-

294 Check List 14 (2)

Figures 2-5. F. hagenii soldiers. 2. Head and pronotum in dorsal view. 3. Oblique view of head. 4: head and pronotum in lateral view. 5.
Lateral view of pronotum. Scale bars: Figs 2-4 = 200 um, Fig. 5 = 100 um.

diers, pseudergates, and immature individuals (larvae and microhabitats occupied by £. hagenii in natural habitats,
white soldiers) (Fig. 7). such as inside a dead hardwood trunk in a Rio de Janeiro
For E. hagenii, there are few data available on the forest (Araujo 1970). This species does not construct

Table 1. Morphometric characters (mm) of Eucryptotermes hagenii soldiers and imagoes from Colonia Benitez (Chaco, Argentina). *New data.

Soldiers (n = 10) Imagoes (n=10)

Morphometric character

Length of head to lateral base of mandibles
Length of head including mandibles*
Length of head to tip of labrum*

Length of head to postclypeus*

Width of head with eyes, maximum

Height of head excluding postmentum
Frontal flangewidth*

Diameter of eye, maximum

Eye to head base distance, minimum*
Diameter of ocellus, maximum

Length of left mandible, maximum

Width of pronotum, maximum

Length of pronotum, maximum

Length of hind tibia

Total body length without wings*

Total body length with wings*

Length of fore wing without scale, maximum*
Width of fore wing, maximum*

Mean (range)
0.99 (0.83-1.05)
1.87 (1.75-2.10)

1.24 (1.13-1.33)
0.99 (0.88-1.15)
1.22 (1.15-1.28)

0.15
0.87
1.14
0.76
0.74
4.44

0.14-0.15
0.80-1.00
1.00-1.28
0.60-0.86
0.64-0.84
3.35-5.36

ee Se es ee

)
)
)
)
)
)

Mean (range)
1.17 (1.06-1.29)

1.46
1.11
1.12
0.73

1.38-1.53
0.90-1.47
1.02-1.22
0.70-0.75

ae aN

)
)
)
)

0.37 (0.34-0.41)
0.15 (0.12-0.18)
0.16 (0.14-0.18)

1.03 (0.99-1.06
0.68 (0.56-0.72
1.05 (0.92-1.13
5.15 (4.20-6.36
9.68 (9.30-10.00)
7.66 (7.40-7.80)
2.11 (2.05-2.20)

)
)
)
)

Godoy et al. | Eucryptotermes hagenii in Chaco, Argentina

295

Figures 6-8. Individuals of E. hagenii and nesting site. 6. Soldier. 7. Alate, soldier and pseudergates. 8. Location of F. hagenii colony inside a
trunk of N. megapotamica living tree. Scale bars = 0.5 mm.

conspicuous arboreal nests like some termitids, and thus,
its detection is difficult and requires exhaustive searching
of standing trees and pieces of dead wood. We found this
species in galleries excavated inside the trunks of living
trees (Fig. 8) and in standing dead wood, as well as in
fallen branches. Native trees having colonies of E. hage-
nii were found to be Nectandra megapotamica (Spreng. )
Mez and E. contortisiliquum (Vell.) Morong. Fallen
branches sheltering £. hagenii were poorly degraded, and
the wood still retained its structure, although some had
high moisture content.

Eucryptotermes, like other “drywood termites”, have
evolutionary characteristics that are considered basal for
termites in regard to their morphology, social organiza-
tion, and nesting type. Nevertheless, the phragmotic head
of soldiers is an advanced or derived condition within
the family Kalotermitidae; with the addition of promi-
nent mandibles, it is developed as a defensive adaptation
to block galleries in the wood and prevent the entry of
predators (Krishna 1961, Scheffrahn et al. 1998).

Nine of the 18 morphometric characters measured
in soldiers and alates of E. hagenii provide new data for
the species; our measurements of the other 6 characters
extend previously known ranges (Constantino 1997). Our
measured values correspond to observations of Constan-
tino (1997), who pointed out that the size of the soldiers
varied considerably. His sample size was small, and

while there was no clear geographical pattern, the largest
soldiers seemed to be in the southern part of the then-
known range, Parana and Santa Catarina states, Brazil.
We cannot corroborate Constantino’s (1997) observations
with our specimens from Chaco; the minimum values of
length and height of head, length of pronotum and length
of hind tibia are smaller than those reported by him.

The previous distributional records of the 2 species of
Eucryptotermes correspond to the Brazilian and Chaco
Subregions within the Neotropical Region. Thus, £.
breviceps is known from a single locality of the Boreal
Brazil Dominion, Roraima Province, whereas FE. hagenii
has been found in the Parana Dominion, Atlantic Prov-
ince. Both species are recorded from native tropical and
subtropical rainforests.

This genus Eucryptotermes and E. hagenii had been
recorded from only 10 places in Brazil (Muller 1873,
Araujo 1977, Constantino 1997, Fontes 1998). Our new
records extend the distribution of the genus Eucryp-
totermes by approximately 980 km west to the Chaco
Dominion, which is characterized by little precipitation
and mesophytic or xerophytic forests (Cabrera and Wil-
link 1973, Morrone 2001). The occurrence of E. hagenii
in this dominion shows that this species has a broader
ecological plasticity than what was previously known.
Our records are also the first for both the genus Eucrypto-
termes and E. hagenii from Argentina. Both the genus and

296

Species were previously thought endemic to Brazil. The
number of termite genera in Argentina is increased to 34
(Silvestri 1903, Torales et al. 1997, 2005, Roisin 2003).

Acknowledgements

We are grateful to Javier Cardeli and Griselda Oria of the
Los Chaguares Natural Reserve for allowing us to con-
duct research in this protected area. Funding was provided
by ANPCyT (PICTO-UNNE 0244) and SGCYT (UNNE).

Authors’ Contributions

All authors collected the field data; MCG identified the
specimens and wrote the manuscript; JMC, GA, and ERL
measured the specimens; CE took the digital and SEM
photographs.

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