ORIGINAL ARTICLE /ARTÍCULO ORIGINAL
A NEW SPECIES OF TEMNOCEPHALA (PLATYHELMINTHES, TEMNOCEPHALIDA)
IN MOUNTAIN CRABS FROM TOLIMA, COLOMBIA
UNA NUEVA ESPECIE DE TEMNOCEPHALA (PLATYHELMINTHES,
TEMNOCEPHALIDA) EN CANGREJOS MONTANOS DE TOLIMA, COLOMBIA
1-2* 3 4
Julian Yessid Arias-Pineda ; Cristina Damborenea & José Joaquín Castro Avellaneda
1Laboratorio de Zoología y Ecología Acuática LAZOEA, Universidad de los Andes, Bogotá, Colombia.
jy.arias10@uniandes.edu.co.
2Grupo de investigación en artrópodos “Kumangui”. Bogotá. Colombia.
3División Zoología Invertebrados, Museo de La Plata, Buenos Aires, Argentina. CONICET. cdambor@fcnym.unlp.edu.ar.
4Departamento de Biología. jjcastro@udistrital.edu.co. Universidad Distrital Francisco José de Caldas, Colombia.
* Corresponding author
Neotropical Helminthology, 2015, 9(2), jul-dec: 253-263.
ABSTRACT
Keywords: Colombia - Ectosymbiont - Neotropical region - South America - Taxonomy -Temnocephala icononcensis n.sp.
Temnocephala icononcensis n.sp. an ectosymbiont on several species of montane
Pseudothelphusidae crabs such as, Hypolobocera bouvieri, Phallangothelphusa dispar and
Strengeriana cajenensis, is described from brooks and small streams in the mountains of Tolima,
Colombia. Eggs and adult specimens were found on the carapace and orbital cavities of the hosts.
The morphology of the species was studied with whole mounted specimens. The most distinctive
character of the new species includes the size, shape and introvert morphology of the cirrus and
the absence of sphincter in the vagina. The finding of this species in Pseudothelphusidae of
Colombia demonstrates the potential diversity of temnocephalans in this country.
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ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043
RESUMEN
Palabras clave: Colombia - Ectosimbiontes - Región Neotropical - América del sur - Taxonomía.
Una nueva especie de Temnocephala Blanchard, 1849, Temnocephala icononcensis n.sp.
ectosimbionte de varias especies de cangrejos montanos de la familia Pseudothelphusidae,
Hypolobocera bouvieri, Phallangothelphusa dispar y Strengeriana cajenensis, encontrado en
quebradas y pequeños arroyos en las montañas del Tolima, Colombia. Huevos y adultos de los
especímenes fueron encontrados en el caparazón y cavidades orbitales de los hospederos. La
morfología de la especie fue estudiada con ejemplares enteros montados. El carácter más
distintivo de la nueva especie incluye el tamaño, la forma y la morfología del introverto de los
cirros y la ausencia de esfínter en la vagina. El hallazgo de esta especie en cangrejos de la familia
Pseudothelphusidae para Colombia demuestra la diversidad potencial de temnocéfalos para este
país.
Colombia, considered as a biodiversity hotspot
area, has 88 Pseudothelphudidae described
species, being 78 of them endemic
(Cumberlidge et al. 2009, 2014). However, no
temnocephalan species associated with
crustaceans are recorded for this country. Only
Temnocephala colombiensis Garcés et al.,
2013, has been described in association with
the snail Pomacea sp. from the Department of
Antoquia, Colombia (Garcés et al., 2013).
During a research work done in Tolima,
Colombia, between 2009 and 2012, several
temnocephalans specimens were found
associated with three pseudothelphusid
species. These individuals were identified as a
new species, which is described in this article.
The new species was compared with the
morphology of other temnocephalan related to
Pseudothelphusidae.
This finding is very important since it is the
first Temnocephala species found living in
association with crabs from Colombia.
Considering the unique characteristics of the
crab fauna in Colombia, this discovery
indicates the high potential diversity of
Temnocephala waiting to be studied.
E i g h t y e i g h t c r a b s b e l o n g i n g t o
Phallangotelphusa dispar (Zimmer, 1912),
Strengeriana cajaensis Campos & Rodriguez,
1 9 9 3 a n d H y p o l o b o c e r a b o u v i e r i
(Rathbum,1898) were collected on brooks and
streams from Cafreria, Hoya Grande, Pie de
Cuesta and Valparaiso, Icononzo Municipality
in Tolima Department (the samples were taken
between 4º06.515 N - 74º35.395 W and
4º06.669 N - 74º35.215 W) (Table 1). The
crabs were manually collected from March
2009 to September 2012. Females and
juveniles crabs were released alive in the
sample site. Crabs were examined in the field
searching for temnocephalans.
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Temnocephala Blanchard, 1849 is a genus of
commensal turbellarians, endemic to the
Neotropical region (Pereira & Cuocolo, 1941;
Cannon, 1993; Damborenea & Cannon, 2001;
Garcés et al., 2013). The first species described
was T. chilensis (Moquin-Tandon, 1846) from
the anomuran crab Aegla laevis (Latreille,
1818) from Chile. Since then around 33
species have been described, associated to
freshwater invertebrates (i.e. Crustacea:
Brachyura, Anomura, Caridea and Astacidea;
Mollusca: Ampullariidae and Neritidae;
Insecta: Belostomatidae, Naucoridae and
Odontoceridae) and turtles (i.e. Chelidae and
Emydidae) (Garcés et al., 2013; Martínez-
Aquino et al., 2014; Seixas et al., 2015).
Brazil, Argentina and Uruguay are the
countries where these species have been
studied more in depth.
The freshwater crab fauna encountered in the
n e o t r o p i c s i n c l u d e s t w o f a m i l i e s
(Trichodactylidae and Pseudothelphusidae)
with approximately 311 species (Cumberlidge
et al., 2014). Despite the great species
diversity, only eight temnocephalan species
have been described associated with seven
species of Trichodactylidae (i.e. Dilocarcinus
pagei Stimpson, 1861, D. septemdentatus
(Herbst, 1783), Sylviocarcinus australis
Magalhães & Türkay, 1996, S. pictus (H. Milne
Edwards, 1853), Trichodactylus fluviatilis
Latreille, 1828, T. panoplus (von Martens,
1869), and T. petropolitanus (Göldi, 1886),
and only five with Pseudothelphusidae
(Hypolobocera henrici Pretzmann, 1978,
Kingsleya ytupora Magalhães, 1986,
Pseudothelphusa tristani Rathbun, 1896, P.
jouyi Rathbun, 1893, and Telphusa sp.)
(Martínez-Aquino et al., 2014).
Pseudothelphusidae has the highest species
diversity in the rivers and streams of the
montains on Colombia and Ecuador.
INTRODUCTION
Arias-Pineda et al.
MATERIALS AND METHODS
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Approximately 50 commensals were
recovered from the coxae and pereipods of
seventeen crabs. The crabs were determined by
specialized key Campos 2005 and deposited in
the collection of arthropods and other
invertebrates CAUD-216. The turbelarians
were fixed at the moment in 70% ethanol and
were transported to the laboratory for posterior
processing. The worms were flattened between
cover and slide, transferred to AFA and kept in
this solution overnight. The AFA solution was
replaced by ethanol 50%, followed by 70%,
and stained overnight with acetocarmine. Then
the Temnocephala were clarified in acidified
ethanol (0.5 ml HCl in ethanol 70%),
dehydrated in ascending series of ethanol
(80%, 90%, and 100%) for 15 min in each,
cleared with a mixture of 1:1 ethanol methyl
salicylate, and later in 100% methyl salicylate.
The specimens were transferred to xylene and
mounted in permount mounted medium
(Fisher Scientific Co., N. J.).
Measurements were taken from fixed and
mounted specimens and expressed in microns
(µm), unless specified otherwise. The mean is
shown, followed by the number of specimens
measured and the size range. The terminology
used for the description of reproductive
structures, were recognized using Cannon´s
(1993) diagnosis. The specimens were
photographed and drawn using the programs
Corel Draw X5 and Photoshop.
The studied material was deposited in the
Colección Artrópodos y otros Invertebrados,
Universidad Distrital Francisco José de Caldas
(CAUD), Bogotá, Colombia and in the Museo
de Hi sto ria Na tur al, Co lec ci ó n d e
Invertebrados, Universidad de los Andes
(ANDES-IN), Colombia.
Family Temnocephalidae Monticelli, 1899
Temnocephala Blanchard, 1849
RESULTADOS
Temnocephala icononcensis n. sp.
(Fig. 1-4)
Specimens examined: 30 specimens of
Temnocephala icononcencis n. sp, collected in
17 montain crabs: Hypolobocera bouvieri,
n=3, Phallangothelphusa dispar, n=8 and
Strengeriana cajaensis, n=6.
Type locality: Colombia, Tolima, Icononzo,
Cafrerias, Las Lajas brook, 4° 06' 51'' N 74° 35'
39'' W, 1,029 meters above sea level; July 2th,
2010; collected by J. Y. Arias-Pineda and R.
Barbosa Cornelio.
Etimology: The species has been named after
the locality were they were collected,
Icononzo, Tolima, Colombia.
Holotype: Specimen stained and mounted in
toto, deposited in CAUD 216 -TEM-0001.
Paratypes: Ten specimens stained and
mounted in toto, deposited in ANDES-IN
(3125), ANDES-IN (3126), ANDES-IN
(3127), ANDES-IN (3128), ANDES-IN
(3129), ANDES-IN (3130), CAUD 216-
TEM-0002, CAUD 216- TEM-0003, CAUD
216- TEM-0004, CAUD 216- TEM-0005,
CAUD 216- TEM-0006.
Host species: Hypolobocera bouvieri
(Rathbum, 1898) CAUD 216 -DEC-0004,
CAUD 216-DEC-0006; Phalangothelphusa
dispar (Zimmer, 1912) CAUD 216-DEC-
0001, CAUD 216-DEC-0002, CAUD 216-
DEC-0003; and Strengeriana cajenensis
Campos & Rodriguez, 1993, CAUD 216-
DEC-0005, CAUD 216-DEC-0007.
Site of infection: Coxopodite of ambulatory
legs, around the eyes and on the ventrolateral
sides of the carapace.
Tipe Host: Phallangothelphusa dispar
(Zimmer, 1912), Colombia, Tolima, Icononzo,
Cafrerias, Quebrada La Fría, 1,029 a 1,258
m.a.s.l., 1 (ancho del caparazón 30.7 mm,
largo del caparazón 28.5 mm). Juny 2th, 2009,
collected by J. Y. Arias-Pineda and R. Barbosa
Cornelio (Fig. 1A-B).
Description
External features. Body ovoid, adult length
Temnocephala icononcensis n.sp., Colombia
Table 1. Samples sites at Icononzo (Tolima, Colombia) and host species. The samples were taken between 4º06.515
N - 74º35.395 W and 4º06.669 N - 74º35.215 W.
Host Localities Altitude Date
Hypolobocera bouvieri
1♂
MUD 046 -DEC-0006
Icononzo, Pie de Cuesta, Quebrada
Mataburros
1150 March 28th
2009
Phallangothelphusa dispar
1♂
MUD 046-DEC-0003
Icononzo, Cafrerias, Quebrada La Fría,
1029 June 2nd 2009
Phallangothelphusa dispar
3♂,3♀
MUD 046 -DEC-
0001
Icononzo, Hoya Grande, Quebrada
Mataburros
1119 August 25th
2009
Phallangothelphusa dispar
3♂,1♀
MUD 046 -DEC-
0002
Icononzo, Cafrerias, Quebrada Las
Lajas
850
October 2009
Strengeriana cajaensis
3♂, 2 ♀
MUD 046 -DEC-
0007
Icononzo, Cafrerias, Quebrada Las
Lajas
1450 April 2nd
2010
Strengeriana cajaensis
5♂, 2 ♀
MUD 046 -DEC-
0005
Icononzo, Pie de Cuesta , Quebrada
Las Lajas
1320 June 2nd 2010
Hypolobocera bouvieri
1♂MUD 046 -DEC-0004
Icononzo, Cafrerias Quebrada La Lajas 1150 September
14th 2012
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without tentacles 1,403.40 μm (n=30; 1000 μm
- 1840 μm), maximum width 804.49 μm (n: 30;
560 μm- 1040 μm). Adhesive disk circular,
diameter 320 μm (n: 30, 220 μm- 400 μm),
occupying the third posterior of the body, with
a small peduncle (Figs. 1C-D, 2A and 3A).
Eyespots red. The excretory syncytial plates
were not observed. Genital pore located in the
middle portion of the body.
Digestive system. A large muscular ovoid
pharynx, width 296.56 μm (n: 30, 200 μm- 450
μm), length 227.58 μm (n: 30; 160 μm- 320
μm), was found in the first third of the body
(Fig. 3C). The intestinal sac is wide and ends at
the level of the middle of the body, not
surpassing it.
Excretory system. A pair of excretory vesicles
very conspicuous, located at the same level of
the mouth. The excretory pores are very
evident (Figs. 2A and 3F). Glands.
Rhabditogen glands formed by numerous
globular, elongated glands, creating clusters of
cells on the lateral sides of the body, extending
from the caudal region of the intestine to the
level of the excretory pore, nor surpassing it
(Figs. 2A and 3A). In the caudal region,
posterior to the intestine sac, adhesive glands
are present. Their ducts are very evident and
converge in the adhesive disk (Fig. 3E). In this
caudal region, in a more central position, two
big cells, paranephrocytes, are observed.
Female reproductive system. Ovary circular,
small, 73.5 μm (n: 30; 56.8μm- 91.3μm) long,
54 μm (n: 30; 37 μm- 67 μm) wide. Vesicular
resorbens rounded, 82.3 μm (n: 30; 40.3 μm-
80.2 μm); its anterior wall contact the intestinal
wall. Seminal receptacles are not observed. A
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Figure 1. Phallangothelphusa dispar and Temnocephala icononcensis n. sp. A-B, dorsal (A) and ventral (B) view of P. dispar,
(scale bars: 25 mm); C-D, specimens of T. icononcensis n. sp. on a pereiopod of a crab (C, scale bar: 1,500 μm) and detail of an
adult (D, scale bar: 1,000 μm); E-F, eggs capsules of T. icononcensis n. sp. deposited on the carapace (E) and on the orbital angle of
P. dispar (F) (scale bars: 350 μm); G, detail of the egg capsules (scale bar: 200 μm).
wide vagina runs to the genital atrium. No
conspicuous vaginal sphincter is observed.
Vitelline glands branched, covering the dorsal
and ventral surface of the intestine. Genital
atrium large and elongated. Genital pore
encircled by abundant small cement glands
(Figs. 2B, 3A-B and D). Eggs claviform, fixed
by a peduncle, 220 y 330 μm long by 30 y 63
μm wide, subpolar filament (Figs. 1E-G).
Opercular plates at small angle respect to
longitudinal axis, the fracture plane is oblique
(Fig. 2D). Male reproductive system. Four
testes, anterior testes 74.02 μm (n = 30, 61.7
μm- 83.98 μm) long, 75.7 μm (n: 30, 61.8 μm -
86.45μm) wide; posterior testis 96 μm (n: 30,
70 μm - 113 μm) long, 94 μm (n: 30, 80 μm –
113.6 μm) wide. The anterior pair of testicles
lateral to the posterior region of the intestine, at
the level of the ovary, posterior testis posterior
to the anterior ones, slightly overlapping.
Seminal vesicle small and elongated (Figs. 2B-
C, 3B-C and 4). Cirrus curved in the third distal
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Arias-Pineda et al.
Figure 2. Temnocephala icononcensisn. sp., holotype. A, adult specimen (scale bar: 250 μm); B, detail of the reproductive system
(scale bar: 100 μm); C, detail of the distal male reproductive system; D, egg and plane of fracture of the operculum, filament,
peduncle (scale bar: 100μm). ad: adhesive disk, at: anterior testes, c: cirrus, dg: adhesive disk glands, ea: excretory ampullae, ga:
genital atrium, gp: genital pore, i: intestinal sac, m: mouth, ov: ovary, pb: prostatic bulb, ph: pharynx, pt: posterior testis, rg:
rhabditogenic glands extending along sides of intestinal sac, sv: seminal vesicle, t: tentacles, v: vitellarium, va: vagina, vr:
vesicula resorbens, arrow: paranephrocyte.
e
ov
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Temnocephala icononcensis n.sp., Colombia
Figure 3. Temnocephala icononcensis n. sp., A, general view of the holotype (scale bar 200: μm); B, detail of the genital system
(scale bar 200: μm); C, detail of the pharynx (scale bar 200: μm); D, cirrus and the gonopore showing the cement glands (scale bar:
200 μm); E, detail of the posterior region, the conducts of the adhesive glands are evident (posterior end to the right) (scale bar: 50
μm); F, excretory ampullae (scale bar: 50 μm).at: anterior testes, c: cirrus, dg: adhesive disk glands, ea: excretory ampullae, m:
mouth, ov: ovary, pb: prostatic bulb, ph: pharynx, pt: posterior testis, , vr: vesicula resorbens.
part, with the concave curvature towards the
intestine; 242.45 μm (n=30; 120 μm- 300 μm)
long, basal wide 75.42 (n: 30, 50 μm - 110 μm),
cirrus introvert 26.25 μm (n: 30, 25 μm - 30
μm) long, with 30-40 rows of spines, each with
approximately 30 spines. The introvert
expands slightly. Some specimens has the
distal part curved (Fig. 4 D) almost at right
angle, while others are almost straight (Fig.
4E). The number of rows of spines can vary
between individuals.
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Arias-Pineda et al.
1941); T. mexicana Vayssiere, 1898 from
Mexico (Lamothe-Argumedo 1968) and T.
peruensis Ibáñez Herrera & Jara, 2003 from
Peru (Ibáñez & Jara 2003). T. lutzi was
originally described in association with
Telphusa sp. (Monticelli 1913). Posteriorly,
this specie was frequently found associated
with trichodactylids crabs in Brazil, Peru and
Uruguay (Pereira & Cuoccolo 1941,
Damborenea 1994, Amato et al. 2005,
Volonterio 2007) and it is discussed by Amato
et al. (2005), that the original host of T. lutzi
might be misidentified. Also, different
concerns about this species have been
mentioned by Volonterio (2007), due to
different morphology interpretations. On the
other hand, the description of T. peruensis,
associated with Hypolobocera henrici, is
DISCUSSION
Only one species of temnocephalan,
Temnocephala colombiensis, is known for
Colombia, and lives associated with Pomacea
sp. (Mollusca, Ampullariidae) (Garcés et al.,
2013). Temnocephala icononcensis n. sp. is the
first temnocephalan species living on
crustaceans for Colombia.
Five temnocephalan species associated with
P s e u d o t h e l p h u s i d a e a r e k n o w n :
Temnocephala costarricensis Lamothe-
Argumedo, 1974 from Costa Rica (Lamothe-
Argumedo 1974); T. kingsleyae Damborenea,
1994 and T. lutzi Monticelli, 1913 from Brazil
(Damborenea 1994, Pereira and Cuocolo,
Figure 4. Cirrus structure of Temnocephala icononcensis n. sp. A-B, Photography from a dissection of a juvenile specimen, cirrus
showing fine spines (scale bar= 50 μm); C, line drawing of cirrus (scale bar= 10 μm); D-E, cirrus of two specimens showing
variation in the curvature (scale bar: 100 μm).
261
insufficient for an accurate identification of the
species.
The cirrus morphology of Temnocephala
icononcensis n. sp. is unique, and it could not
be compared with other Temnocephala species
from Pseudothelphusidae. The temnocephalan
species with a comparable introvert with one
of the new species is Temnocephala chilensis,
species associated with several species of
Aegla sp. (Crustacea, Aeglidae) and
Samatacus sp. and Parastacus sp. (Crustacea,
Parastacidae). T. chilensis has an introvert
more expanded than the new species, and the
spines are stronger than the ones in the new
species; however, the size and the shape of the
cirrus in both species are different (Dioni,
1967, 1972; Damborenea, 1991, 1992). The
cirrus of T. chilensis is shorter (149.59 um long
and 62.91 basal width), after Damborenea
(1992) and the cirrus never has a curvature so
pronounced as observed in the new species.
Other feature share by both species is the lack
of a vaginal sphincter.
Among the limitations of this study, the first
was the difficult access to the study area, and
difficult collection of specimens by the
topography of the area. The host and
temnocephalos populations vary with respect
to the season, being in some months more than
others.
The cirrus morphology is the most relevant
feature for the identification of temnocephalid
species. Other relevant structures for species
diagnosis are the presence and development of
vaginal sphincter, the general morphology of
the genital organs, and the pharynx. All these
characters could be studied and described. Size
and morphology of the excretory syncytial
plates are features that allow comparison
between the temnocephalan species.
Unfortunately, the syncytial plates are not
known for temnocephalan species from
Pseudothelphusidae, and they could not be
studied for the new species. Future studies of
Amato, JFR, Amato, SB & Seixas, SA. 2005.
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