Tugas Ringkasan PDF

Emerging Topics in Life Sciences (2017) 1 651–657
https://doi.org/10.1042/ETLS20170111
Trematoda (flukes)
Rafael Toledo1 and Bernard Fried2
1
Departamento de Parasitología, Facultad de Farmacia, Universidad de Valencia, Av. Vicente Andrés Estellés s/n, 46100 Burjassot, Valencia, Spain; 2Department of Biology,
Lafayette College, Easton, PA 18042, U.S.A.
Correspondence: Rafael Toledo (rafael.toledo@uv.es)
The class Trematoda is the largest group of Platyhelminths and includes two subclasses:
Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Aspidogastrea and Digenea. Trematodes, and particularly Digeneans, is a large group of
organisms with significant medical and veterinary interest. Over 100 species of digenetic
trematodes have been reported infecting humans. Although the significant mortality and
morbidity that some of these infections cause, they are among the most neglected trop-
ical diseases. Apart from their impact in public and animal health, the Digenea constitutes
an intriguing group of organisms that has a vast interest in experimental biology.
Systematics and taxonomy of this group constitute a challenge for biologists in relation
to the difficulty entailed in the establishment of phylogenetic relationships between tre-
matodes and the determination of valid diagnostic features. Moreover, their complex life
cycles, using at least two hosts and alternating free-living and parasitic stages or sexual
and asexual multiplication, constitute a paradigm of how organisms can evolve to
become adapted to different biotic and abiotic environments to enhance survival. In this
review, we briefly summarize the major features of trematodes in relation to both bio-
logical and medical areas.
Introduction
The class Trematoda constitutes a large group of Platyhelminthes parasites that is divided into two sub-
classes: the Aspidogastrea and the Digenea. The Aspidogastrea, also called aspidobothrea, is a small
group comprising 4 families, 12 genera and ∼80 species. In general, they are parasites of freshwater
and marine mollusks, and cartilaginous or bony fishes and turtles may serve as facultative or obligatory
hosts. Commonly, Aspidogastrea are external parasites of these hosts and have a single-generation life
cycles lacking asexual reproduction. Adults are conspicuous for having a large ventral disc often
divided into several compartments. There are no species of Aspidogastrea with medical or economic
importance. However, further information can be found in the works by Prof. Klaus Rhode [1,2].
In contrast, the subclass Digenea is a markedly larger group and has a significant medical, veterin-
ary and economic importance. Digeneans, also called digenetic trematodes or flukes, comprises more
than 18 000 species that are obligatory parasitic in invertebrate intermediate and vertebrate definitive
host [3]. Owing to their greater importance, this review will be focused on the subclass Digenea. Most
digenean species inhabit the intestine as adults, but several species colonize other habitats within the
vertebrate host. Digenetic trematode infections occur worldwide and over 100 species have been
recorded infecting humans causing serious problems in them. For example, it is estimated that almost
300 million people are currently infected with Schistosoma spp. causing ∼6,000 deaths annually [4].
Moreover, a large list of digenetic trematodes actually parasitize animals both of veterinary interest
and wildlife.
Apart from their medical and veterinary importance, digenetic trematodes have a vast interest in
Received: 11 September 2017 experimental biology due to many characteristics of these parasitic organisms: (i) the paradigmatic
Revised: 17 October 2017 systematics of this group of parasites in relation to the morphological similarity between members of
Accepted: 20 October 2017
different taxa and the inadequate or poor specific diagnosis (or both) of several newly established
Version of Record published: taxa. Furthermore, the absence or loss of type material for several taxa further complicates the situ-
22 December 2017 ation. As a result, this group has been characterized by a long history of synonymies and inadequate
© 2017 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society and the Royal Society of Biology 651
diagnosis. (ii) The complexity of the life cycles that has led to the development of an important number of
adaptive strategies to enhance parasite survival and transmission converts these organisms to an excellent
model to study the evolutionary adaptation of an organism to an environment.
Taxonomy and functional morphology

The taxonomy of the Digenea has long been a challenge due to the difficulties in establishing relationships
and diagnostic features, even for the higher taxa [3]. In fact, more than 2500 genera have been historically
accepted within this group. Relatively recently, an important effort was done to clarify the systematics of this
group by Dr Gibson and colleagues. This work was reflected in a series of three volumes (Keys to the
Trematodes) that exhaustively review the taxonomy of these parasites [5–7]. This relevant piece of work pro-
vides historical background and novel approaches for the systematics and taxonomy at the higher taxa and a
re-appraisal of the generic diagnosis. In fact, only 148 families with 1577 genera were considered valid. In

spite of this detailed work, there are still several gaps in our knowledge on this topic and further effort is
required. In this context, molecular analysis may be key to achieve an improved picture of the digenetic
taxonomy.
Figures 1 and 2 show details on the morphology of adult Digenean Trematodes. Digenetic trematodes have a
dorsoventrally flattened body, bilateral symmetry and a definite anterior end. They are leaf-shaped and may
range in size from a few millimeters to ∼8 cm (i.e. Fasciolopsis buski). The form and function of major organ
systems of the digenetic trematodes has been recently reviewed by Peoples and Fried [8].
The body surface of flukes comprises a syncytial tegument. The epidermis comprises a distal anucleate cell
layer. Cell bodies containing the nuclei lie beneath a superficial layer of muscles, connected to the cytoplasm
by several channels [8]. Moreover, tegument can be covered by spines and also molecules released by the
worm can be found [9]. Adult flukes possess an oral sucker around the mouth and a ventral sucker or acet-
abulum that can be used for the attachment to the host tissues (Figures 1 and 2). Apart from adult stages, life
cycle of trematodes may include up to six larval stages known as egg, miracidium, sporocyst, redia, cercariae
and metacercaria. In some of these larval stages, some particularities can be found. The miracidium is covered
by ciliated epithelial cells that are shed during transformation to either sporocyst or redia. Early embryos of
cercariae are covered with a primary epidermis, and the definitive epithelium is formed underneath of this
primary epidermis. The wall of the metacercarial cyst is formed by a material released from the cystogenic
cells found within the parenchyma of the cercariae. Recently, it has been shown that several trematodes are
able to produce secretomes that are actively secreted containing molecules of interest in communication with
the host [10]. The nervous system consists of longitudinal cords connected by transverse ring commissures
[8]. Moreover, flukes possess a variety of sensory adaptations including photosensory, chemosensory and
mechanosensory [11,12]. Commonly, muscles are circular lying beneath the basal lamina of the tegument in
lateral areas.
Internal organs are embedded in connective tissue parenchyma. Flukes have an alimentary canal including
pharynx and esophagus. The intestine is commonly a branched tube. The main branches may end blindly or
open into an excretory vesicle. The excretory vesicle also accepts the two main lateral collecting ducts of the
excretory system which is of a protonephridial type with ciliated flame cells. These cilia facilitate excretion of
fluids through an excretory pore, commonly located at the posterior end [8].
Most trematodes (excluding members of the family Schistosomatidae) are hermaphroditic having each indi-
vidual with both male and female reproductive system. Male organs include two testes with accessory glands
and ducts leading to a cirrus that extends into a common genital atrium (Figure 2c). The female system consists
of an ovary including seminal receptacle and vitellaria that connect with the oviduct and it expands into an
ootype. The uterus opens into the genital pore into the genital atrium. Both self- and cross-fertilization may
occur depending on the species. In the case of schistosomatids, the structure of the reproductive systems is
similar, but these flukes are dioecius and there are separate male and female individuals.
Biology of trematodes
One of the most relevant features of flukes is the complexity of their life cycle which are characterized by: (i)
alternation of several generations known as the adult, egg, miracidium, sporocyst, redia, cercariae and metacer-
caria; (ii) alternation of sexual and asexual multiplication; (iii) an intensive asexual multiplication enhancing
the transmission of the parasite and (iv) inclusion of two or three hosts, one invertebrate intermediate host
(usually a mollusk) in which asexual multiplication occurs and a vertebrate definitive host in which adult
652 © 2017 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society and the Royal Society of Biology

Figure 1. Organography of an adult Trematode.
Generalized diagram of the organography of an adult digenetic trematode.
worms reproduce sexually. All these characteristics enable the trematodes to be an excellent model to study the
parasite–host interactions in both invertebrate and vertebrate organisms [13,14].
Although each species has a unique life history, a general cycle is as follows. Adult worms produce eggs in
the vertebrate definitive host. Eggs are passed in the feces, urine or sputum, depending on the location of adult
worms, to reach the external environment. The eggs are either eaten by a mollusk and the miracidia hatch, and
invade this first intermediate host, or hatch in the external environment allowing the miracidium to locate and
penetrate the appropriate molluskan host. Within the mollusk first intermediate host, asexual multiplication
occurs, which is critical to enhance the transmission of the parasite. The miracidia produce sporocysts that
contain germ cells which will give rise to a large number of the subsequent generations of larval stages.

Figure 2. Microphotographs of Trematodes with details on several organs.
(a) Adult specimen of a digenetic trematode (Echinoparyphium recurvatum) stained with borax carmine (scale bar: 1 mm).
(b) SEM microphotograph of an adult digenetic trematode (E. caproni) showing details of the mouth, oral sucker and ventral
sucker (scale bar: 100 mm). (c) SEM microphotograph of an adult digenetic trematode (E. caproni) showing details of the cirrus
(scale bar: 50 mm).
Sporocysts are asexual sacs containing a large number of germ cells each of which produces either a subsequent
generation of sporocysts or rediae. Rediae are similar to sporocysts, but have a mouth and pharynx to feed dir-
ectly on the host tissues. Rediae share the ability of sporocysts to produce a large number of the subsequent
generations and, ultimately, to cercariae that is the last stage produced in the mollusk and responsible for the
transmission to the next host in the life cycle. Although this ability of asexual multiplication may imply serious
damage for the molluskan host, it enhances the transmission of the parasite in spite of the difficulty entailed in
the encounter between miracidium and first intermediate host and the hostile environment by ensuring the
generation of a large number of the next infective, larval stage infective, from a single miracidium. For example,
it has been shown that an individual miracidium [15] of Echinostoma caproni infecting Biomphalaria glabrata
resulted in the production of almost 1500 cercariae during the first month of cercarial emission [8]. Cercaria,
the ultimate stage of asexual multiplication, is a tailed free-living stage that is released from the mollusk. In
some species (i.e. blood flukes), cercariae enter the vertebrate definitive host where they mature to adult worms.
In other species, cercariae penetrate, or are eaten by, the second intermediate host and encyst as metacercariae,
and the definitive host became infected after ingestion of the second intermediate host, harboring the metacer-
caria. Finally, in other species (i.e. Fasciola hepatica), cercariae swim in an aquatic environment to reach the
aquatic plants where they attach as metacercariae. The vertebrate host becomes infected by eating the
metacercariae.
Trematodes of medical interest

As mentioned above, more than 100 species of digenetic trematodes have been reported infecting humans.
Although most of them are irrelevant since they do not cause symptoms or are mild and transient, there are
many flukes that constitute an important public health problem. Species with medical interest are commonly
classified on the basis of their location in the human host: blood flukes (Schistosoma spp.), liver flukes (F. hep-
atica, F. gigantica, Clonorchis sinensis and Opisthorchis spp.), lung flukes (Paragonimus spp.) and intestinal
flukes. According to their mode of infection and the subsequent measures of control, these parasitic diseases
also are classified into blood flukes, in which cercariae actively penetrate into the human skin in aquatic envir-
onments, and food-borne trematode infections, including liver, lung and intestinal flukes, in which humans
become infected by ingestion of foods harboring the metacercariae. Although the considerable public health
impact and the emerging nature of food-borne trematodiases [4,16,17], they are among the most neglected of
the so-called neglected tropical diseases and it was recognized by WHO [18]. It should be noted that the
neglected tropical diseases are found predominantly in the world’s poorest populations in low-income
countries, and where these diseases are common, they exacerbate poverty. Major features of these parasitic
infections have been recently compiled in the book edited by Toledo and Fried [19].
Blood flukes cause schistosomiasis or bilharziasis that is one of the most important parasitic diseases of
humans causing significant morbidity and mortality on several continents. Schistosomiasis is transmitted in
over 70 countries, throughout a wide belt of the tropics and subtropics [20,21]. The three major species causing
schistosomiasis are Schistosoma haematobium, S. mansoni and S. japonicum. It is difficult to determine the
number of schistosomiasis cases, but it has been estimated that ∼300 million people were infected in 2013
worldwide and ∼280,000 annual deaths occur [4,22]. Although the number of deaths caused by schistosomiasis
decreased from 1990 to 2013, the number of active infections increased 31% in the same period which is evi-
dence for the need to implement campaigns of control [4]. In addition to active infections, it is estimated that
almost 800 million people worldwide are at risk of infection [23].
It is estimated that ∼80 million people worldwide were infected with any species of food-borne trematodes

in 2013, showing a growing trend from 1990 (∼51% greater number of cases in 2013) [4]. However, these data
are controversial, since other authors [24] reported an estimation of ∼56 million people using a systematic
review and meta-analysis.
Fasciola spp. causes an important liver disease called fascioliasis that is an important medical and veterinary
problem of worldwide distribution. The estimates on the incidence of human fasciolasis are confusing and
range from 2.4 to 17 million people [25]. Opisthorchiasis, caused by Opisthorchis spp., and clonorchiasis,
caused by C. sinensis, are the major fish-borne flukes [26]. It is estimated that ∼24 million people are currently
infected [24]. Although symptoms of these diseases are commonly mild, cholangiocarcinoma is the most
serious complication. Opisthorchis viverrini is classified by the International Agency for Research on Cancer
(IARC) as definitely carcinogenic (class 1) and C. sinensis as a probable carcinogen (class 2A) [27].
Cholangiocarcinoma is a malignant tumor starting in the epithelium of the intrahepatic biliary tree and might
invade the sinusoids of the liver parenchyma [26]. The prognosis of this tumor is extremely poor [27]. Lung
infections or paragonimiasis are caused by Paragonimus spp. There are ∼15 species of Paragonimus known to
infect humans. P. westermani is the most common worldwide, while P. heterotremus is the etiological agent of
human paragonimiasis in PR China, Laos PDR, Vietnam and Thailand [28]. About 23 million people are
infected with lung flukes and almost 300 million people are estimated to be at risk of infection [24,28]. The
intestinal food-borne trematodes category constitutes a large assemblage of species that induce parasitic zoo-
noses. Collectively, these parasites affect the health and economy in a big way in developing countries of the
tropics and subtropics in Asia, Africa, Europe and the U.S.A. A total of 70 species (14 families and 36 genera)
of intestinal flukes have been isolated from humans [29–31]. About 7 million people were estimated to be
infected in 2012 [24].
Another point of interest of human trematode infections is related to the treatment of these diseases.
Praziquantel is the drug of choice, though other drugs such as triclabendazole or tribendimidine also can be
used. In general, all of them provide satisfactory results, but there are several concerns in relation to factors that
include: (i) the limited number of available drugs; (ii) the side effects of these drugs; (iii) the commercial avail-
ability of these drugs is limited and is authorized only for human use in particular countries and (iv) there is
increasing evidence in relation to the development of resistance against these drugs [32]. Thus, further efforts
for the developments of new active compounds are required. In this context, several promising synthetic and
natural products are being studied [30].
Conclusion
Trematodes, and particularly digenetic trematodes, constitute an amazing group of parasites for both their
medical and veterinary importance and their interest in more basic biological sciences. Trematode infections
have been considered for a long time as minor diseases confined to low-income countries and their importance
has been underestimated. However, their frequency is greater than previously considered and some of them
have relevant mortality and morbidity. Moreover, these diseases are re-emerging worldwide, including in devel-
oped countries in relation to the ‘globalized world’. Although considerable efforts are being made to control
these diseases, the emergent nature of them requires new epidemiological and pharmacological approaches.
Additionally, trematodes constitute an intriguing group of parasites in relation to the taxonomy and other
factors such as their complex life cycles and the evolutionary adaptations of these organisms to guarantee sur-
vival in different biological environments.
Summary
• The class Trematoda includes a large number of species of medical and veterinary interest.
• Further studies on the systematics and taxonomy of these organisms are required.
• This constitutes an intriguing group of organisms in relation to their complex life cycles and
the biological adaptations to enhance their survival.

Funding
This work was supported by the Projects PROMETEO2014-083 Fase II from Generalitat Valenciana,
BFU2016-75639-P, from the Ministerio de Economía y Competitividad, and No. RD12/0018/0013, Red de
Investigación Cooperativa en Enfermedades Tropicales — RICET, IV National Programe of I+D+I 2008-2011,
ISCIII — Subdirección General de Redes y Centros de Investigación Cooperativa and FEDER from the Ministerio
de Sanidad y Consumo.
Competing Interests
The Authors declare that there are no competing interests associated with the manuscript.
References
1 Rhode, K. (2001) The Aspidogastrea: an archaic group of Platyhelminthes. In Interrelationships of the Platyhelminthes (Littlewood, D.T.J. and Bray, R.A.,
eds), pp. 159–167. Taylor & Francis, London
2 Rhode, K. (2002) Subclass Aspidogastrea Faust & Tang, 1936. In Keys to the Trematoda (Gibson, D.I., Jones, A. and Bray, R.A., eds), vol. 1, pp. 5–14.
CAB International, Wallingford
3 Kostadinova, A. and Pérez-del-Olmo, A. (2014) The systematics of the Trematoda. Adv. Exp. Med. Biol. 766, 21–44 https://doi.org/10.1007/
978-1-4939-0915-5_2
4 Herricks, J.R., Hotez, P.J., Wanga, V., Coffeng, L.E., Haagsma, J.A., Basáñez, M.-G. et al. (2017) The global burden of disease study 2013: what does
it mean for the NTDs? PLoS Negl. Trop. Dis. 11, e0005424 https://doi.org/10.1371/journal.pntd.0005424
5 Gibson, D.I., Jones, A. and Bray, R.A. (eds) (2002) Keys to the Trematoda, vol. 1. CAB International, Wallingford
6 Jones, A., Bray, R.A. and Gibson, D.I. (eds) (2005) Keys to the Trematoda, vol. 2. CAB International, Wallingford
7 Bray, R.A., Gibson, D.I. and Jones, A. (eds) (2008) Keys to the Trematoda, vol. 3. CAB International, Wallingford
8 Peoples, R.C. and Fried, B. (2014) Form and function in the Digenea. Adv. Exp. Med. Biol. 766, 3–20 https://doi.org/10.1007/978-1-4939-0915-5_1
9 Cortés, A., Sotillo, J., Muñoz-Antolí, C., Molina-Durán, J., Esteban, J.G. and Toledo, R. (2017) Antibody trapping: a novel mechanism of parasite
immune evasion by the trematode Echinostoma caproni. PLoS Negl. Trop. Dis. 11, e0005773 https://doi.org/10.1371/journal.pntd.0005773
10 Marcilla, A., Trelis, M., Cortés, A., Sotillo, J., Cantalapiedra, F., Minguez, M.T. et al. (2012) Extracellular vesicles from parasitic helminths contain
specific excretory/secretory proteins and are internalized in intestinal host cells. PLoS ONE 7, e45974 https://doi.org/10.1371/journal.pone.0045974
11 Abdul-Salam, J. and Sreelatha, B.S. (2004) Description and surface topography of the cercaria of Austrobilharzia sp. (Digenea: Schistosomatidae).
Parasitol. Int. 53, 11–21 https://doi.org/10.1016/j.parint.2003.10.001
12 Filippi, J.-J., Quilichini, Y., Foata, J. and Marchand, B. (2012) Topography and ultrastructure of the tegument of Lecithochirium musculus (Digenea:
Hemiuridae), a parasite of the European eel Anguilla (Osteichthyes: Anguillidae). J. Morphol. 273, 361–370 https://doi.org/10.1002/jmor.11032
13 Toledo, R. and Fried, B. (2005) Echinostomes as experimental models for interactions between adult parasites and vertebrate hosts. Trends Parasitol.
21, 251–254 https://doi.org/10.1016/j.pt.2005.04.006
14 Toledo, R., Muñoz-Antoli, C. and Fried, B. (2007) The use of echinostomes to study host–parasite relationships between larval trematodes and
invertebrate and cold-blooded vertebrate hosts. Parasitol. Res. 100, 1177–1185 https://doi.org/10.1007/s00436-007-0470-6
15 Toledo, R., Muñoz-Antoli, C. and Esteban, J.G. (1999) Production and chronobiology of emergence of the cercariae of Euparyphium albuferensis
(Trematoda: Echinostomatidae). J. Parasitol. 85, 263–267 https://doi.org/10.2307/3285630
16 Engels, D. and Savioli, L. (2006) Reconsidering the underestimated burden caused by neglected tropical diseases. Trends Parasitol. 22, 363–366
https://doi.org/10.1016/j.pt.2006.06.004
17 Hotez, P. and Aksoy, S. (2017) PLOS neglected tropical diseases: ten years of progress in neglected tropical disease control and elimination … more or
less. PLoS Negl. Trop. Dis. 11, e0005355 https://doi.org/10.1371/journal.pntd.0005355
18 World Health Organization. Neglected tropical diseases [Internet]. Available from: http://www.who.int/neglected_diseases/diseases/en
19 Toledo, R. and Fried, B. (eds) (2014) Digenetic Trematodes. Springer, New York
20 World Health Organization. Schistosomiasis facts sheet [Internet]. [Updated January 2017]. Available from: http://www.who.int/mediacentre/factsheets/
fs115/en
21 Lewis, F.A. and Tucker, M.S. (2014) Schistosomiasis. Adv. Exp. Med. Biol. 766, 47–75 https://doi.org/10.1007/978-1-4939-0915-5_3
22 van der Werf, M.J., de Vlas, S.J., Brooker, S., Looman, C.W.N., Nagelkerke, N.J.D., Habbema, J.D. et al. (2003) Quantification of clinical morbidity
associated with schistosome infection in sub-Saharan Africa. Acta Trop. 86, 125–139 https://doi.org/10.1016/S0001-706X(03)00029-9
23 Steinmann, P., Keiser, J., Bos, R., Tanner, M. and Utzinger, J. (2006) Schistosomiasis and water resources development: systematic review,
meta-analysis, and estimates of people at risk. Lancet Infect. Dis. 6, 411–425 https://doi.org/10.1016/S1473-3099(06)70521-7
24 Fürst, T., Keiser, J. and Utzinger, J. (2012) Global burden of human food-borne trematodiasis: a systematic review and meta-analysis. Lancet Infect. Dis.
12, 210–221 https://doi.org/10.1016/S1473-3099(11)70294-8
25 Mas-Coma, S., Valero, M.A. and Bargues, M.D. (0000) Fascioliasis. Adv. Exp. Med. Biol. 766, 77–114 https://doi.org/10.1007/978-1-4939-0915-5_4
26 Saijuntha, W., Sithithaworn, P., Kaitsopit, N., Andrews, R.H. and Petney, T.N. (2014) Liver flukes: Clonorchis and Opisthorchis. Adv. Exp. Med. Biol. 766,
153–199 https://doi.org/10.1007/978-1-4939-0915-5_6
27 Sripa, B. and Pairojkul, C. (2008) Cholangiocarcinoma: lessons from Thailand. Curr. Opin. Gastroenterol. 24, 349–356 https://doi.org/10.1097/MOG.
0b013e3282fbf9b3
28 Blair, D. (2014) Paragonimiasis. Adv. Exp. Med. Biol. 766, 115–152 https://doi.org/10.1007/978-1-4939-0915-5_5

29 Toledo, R., Bernal, M.D. and Marcilla, A. (2011) Proteomics of foodborne trematodes. J. Proteomics 74, 1485–1503 https://doi.org/10.1016/j.jprot.
2011.03.029
30 Toledo, R., Esteban, J.G. and Fried, B. (2012) Current status of food-borne trematode infections. Eur. J. Clin. Microbiol. Infect. Dis. 31, 1705–1718
https://doi.org/10.1007/s10096-011-1515-4
31 Toledo, R., Muñoz-Antoli, C. and Esteban, J.G. (2014) Intestinal trematode infections. Adv. Exp. Med. Biol. 766, 201–240 https://doi.org/10.1007/
978-1-4939-0915-5_7
32 Keiser, J. and Utzinger, J. (2004) Chemotherapy for major food-borne trematodes: a review. Expert Opin. Pharmacother. 5, 1711–1726 https://doi.org/
10.1517/14656566.5.8.1711

Tugas Ringkasan PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Tugas Ringkasan PDF

Uploaded by

Copyright:

Available Formats

Emerging Topics in Life Sciences (2017) 1 651–657

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Taxonomy and functional morphology

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Trematodes of medical interest

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

Downloaded from https://portlandpress.com/emergtoplifesci/article-pdf/1/6/651/484031/etls-2017-0111c.pdf by guest on 20 May 2020

You might also like