Review Article |
Corresponding author: Constanza Vásquez-Doorman ( rdoorman@uchile.cl ) Academic editor: Piter Boll
© 2022 Constanza Vásquez-Doorman, Javiera Escobedo, Miguel L. Allende.
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.
Citation:
Vásquez-Doorman C, Escobedo J, Allende ML (2022) Current state of knowledge on freshwater planarians (Platyhelminthes, Tricladida, Dugesiidae) from Chile. Neotropical Biology and Conservation 17(3): 185-203. https://doi.org/10.3897/neotropical.17.e82779
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The unique geography of Chile encompasses a wide diversity of ecosystems and a rich biodiversity. However, the platyhelminth fauna has been poorly studied. The aim of this work is to compile the historical record of freshwater planarians described for this country. We accessed worldwide databases and published articles to provide a comprehensive review of their discovery history, morphological characteristics and their localities. Freshwater planarians have been collected mainly in central and southern Chile, while in the northern region a single species has been described. The discovery of new species of freshwater triclads has the potential to reveal novel animal models to study regeneration and/or biological adaptations, as some species are suitable for culture in the laboratory. We discuss the many reasons why further research is needed for this animal group, which should include genomic and molecular genetic studies.
biodiversity, dugesiids, neotropical region, triclads biodiversity, dugesiids, neotropical region, triclads
The phylum Platyhelminthes comprehends a diverse group of free-living and parasitic flatworms. Free-living Tricladida flatworms are divided into three suborders based on their habitat: Maricola (marine), Cavernicola (cave), and Continenticola (freshwater and terrestrial) (
The reproductive system’s complexity and its diversity makes it ideal for taxonomic classification as a diagnostic character because it offers numerous characters for taxonomic and systematic studies. Sexual triclads are hermaphrodites, each animal has male and female reproductive organs and mate by cross-fertilization. Testes appear as discrete follicular structures distributed ventrally or dorsally along the body, and a pair of ovaries is usually located closely posterior to the brain. Other anatomical structures to consider are the vitellarian follicles or yolk glands distributed along the body and that generally occupy the entire dorsoventral space, as well as, the shape of the penis papilla, muscle types, routes of ovovitelline and seminal ducts, glands, and characteristics of the copulatory bursa.
For many years, the freshwater planarian fauna in Chile was known only by the studies of Borelli and Böhmig in the early 1900s (
To search on the WoRMS database, we selected the distribution tab and chose “Chile (Nation)” under geounit. Then, we limited taxa to “Dugesiidae Ball, 1974” and asked to include all records, unchecking “extant only” and “marine taxa”. To search within the TTD (
The map was generated using QGIS Hannover version 3.16 incorporating the coordinates as delimited text layers (.csv) for each species. Then, we edited the images on Adobe Photoshop 2021 to add location numbers. Planarian drawings and copulatory apparatus sketches were recreated using Adobe Illustrator and Photoshop 2021.
The freshwater family Dugesiidae was coined by
The Italian zoologist Alfredo Borelli described species collected by Enrico Festa, providing the first description of two species from Ecuador and Argentina, but also present in Chile: Planaria festae (G. festae) and Planaria patagonica (R. patagonicus) (
Overall, the described species occur in central (30–40°S) and southern (50–55°S) Chile (Fig.
Localities of freshwater planarian species from Chile. Indicating locality, coordinates, spot map number, collector and/or year of sampling and associated reference for each species.
Sp. | Locality (Chile) | Coordinates | Spot | Collector | Reference |
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Dugesia Girard, 1850 | |||||
D. ambigua | |||||
A mountain lake, Punta Arenas, Magallanes | 53°09'50"S, 70°55'02"W | 22 | J. Michaelsen, 1893 |
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D. similis | |||||
A brook, Salto, Valparaíso | 33°02'56"S, 71°30'42"W | 9 | J. Michaelsen, 1893 |
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Girardia Ball, 1974 | |||||
G. canai | |||||
Reñaca Estuary, Concón, Valparaíso | 32°58'23"S, 71°32'38"W | 6 | Unknown |
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Mapocho River, Talagante, Metropolitana | 33°40'00"S, 70°56'00"W | 11 | P. Sánchez, 1986 |
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G. chilla | |||||
Cadillos and Cajon Brooks, Zapallar, Valparaíso | 32°33'04"S, 71°27'38"W | 2 | Unknown |
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El Tigre Brook, Zapallar, Valparaíso | 32°34'20"S, 71°26'17"W | 3 | Unknown |
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Huallilemu Estuary, Punta de Tralca, Valparaíso | 33°26'13"S, 71°41'29"W | 10 | P. Sánchez |
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Near Pucatrihue Beach, Osorno, Los Lagos | 40°41'00"S, 73°32'00"W ** | 14 | 1991 |
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Pichi Pilluco Estuary, Llanquihue, Los Lagos | 41°27'00"S, 72°55'00"W * | 15 | 1948 |
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A stream, Ancud, Chiloé Island, Los Lagos | 41°51'50"S, 73°49'20"W * | 16 | 1948 |
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A stream, Cancha Ski, Punta Arenas, Magallanes | 53°11'00"S, 70°58'00"W * | 23 | 1949 |
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G. festae | |||||
El Tigre Brook, Zapallar, Valparaíso | 32°34'20"S, 71°26'17"W | 3 | Unknown |
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Catapilco Estuary, La Laguna, Valparaíso | 32°37'36"S, 71°24'58"W | 4 | Unknown |
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El Cobre Estuary, Nogales, Valparaíso | 32°40'17"S, 71°12'56"W | 5 | Unknown |
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Reñaca Estuary, Concón, Valparaíso | 32°58'23"S, 71°32'38"W | 6 | Unknown |
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Limache Estuary, Valparaíso | 33°01'28"S, 71°16'23"W | 7 | Unknown |
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A ditch, Peñablanca, Quilpué, Valparaíso | 33°02'14"S, 71°21'10"W | 8 | J. Michaelsen, 1893 |
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Huallilemu Estuary, Punta de Tralca, Valparaíso | 33°26'13"S, 71°41'29"W | 10 | Unknown |
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Mapocho River, Talagante, Metropolitana | 33°40'00"S, 70°56'00"W | 11 | P. Sánchez |
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Mapocho River, Talagante, Metropolitana | 33°40'00"S, 70°56'00"W * | 11 | M. Sánchez |
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Concepción, Biobío | 36°49'09"S, 73°02'40"W | 12 | I. Hermosilla, T. Cekalocic, 1976 |
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A flow, Valdivia, Los Ríos | 39°49'02"S 73°14'33"W | 13 | J. Michaelsen, 1893 |
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G. rincona | |||||
Elqui River, Vicuña, Coquimbo | 30°02'00"S, 70°42'00"W *** | 1 | 1949 |
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Romankenkius Ball, 1974 | |||||
R. patagonicus | |||||
Stream by Pehoe Lake, Torres del Paine, Magallanes | 51°04'00"S, 73°14'00"W * | 17 | M. Haga, 1966 |
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Pehoe Lake, Paine, Magallanes | 51°04'00"S, 73°14'00"W | 17 | S. Inoue, 1971 |
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Serrano River, del Toro Lake, Magallanes | 51°11'00"S, 73°12'00"W * | 18 | S. Inoue, 1971 |
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Serrano River, near its confluence with the Grey River, Magallanes | 51°14'00"S, 73°13'00"W * | 19 | S. Inoue, 1971 |
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Stream by del Toro Lake with Porteño Lake, Magallanes | 51°19'00"S, 72°59'00"W * | 20 | S. Inoue, 1971 |
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de las Minas River, Punta Arenas, Magallanes | 53°08'00"S, 70°51'00"W * | 21 | M. Haga, 1966 |
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Daly Estuary, Grande Island Tierra del Fuego, Magallanes | 53°33'38"S, 68°42'27"W **** | 24 | M. Haga, 1966 |
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Picton Island, Cabo de Hornos, Magallanes | 54°10'00"S, 71°20'00"W | 25 | J. Michaelsen, 1892 |
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Aracena Island, Cockburn Channel, Magallanes | 55°02'19"S, 66°55'33"W | 26 | 1932 |
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Distribution of freshwater planarians described from Chile and their known distribution in Latin America. We have listed the records (spots 1–26) with the corresponding coordinates in Table
Because most Chilean specimens were collected many years ago, only a few of them have been deposited in international museums. For example, G. festae was deposited in the American Museum of Natural History (
We have summarized the external aspect and some morphological characteristics of each species in Fig.
Summary of morphological characteristics of freshwater triclads from Chile. 1 = number of studied specimens, 2 = animal size in mm (length × width), 3 = presence of mature sexual organs, 4 = testes distribution along the dorsoventral axis, 5 = signs of asexual reproduction, 6 = pharynx pigmentation, ? = information not provided.
1 | 2 | 3 | 4 | 5 | Dorsal coloration | Ventral coloration | 6 |
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D. ambigua | |||||||
? | 7 × 3 | No | D | ? | Deep brown, blackish or dirty olive-green; whitish or yellowish on the margins and midline. | Whitish or yellowish, occasional brownish spots. | ? |
D. similis | |||||||
many | 1.5–7 × 1.5–2.5 | No | ? | Yes | Isolated or confluent dark-brown, almost blackish spots covering the yellow pale background. | Evenly colored light gray or blackish spotted. | ? |
G. canai | |||||||
16 | 9–10 × 2–4 | Yes | V | ? | Light brown, with dark brown spots that might form a median stripe. | ? | Yes |
G. chilla | |||||||
50+ | 9–25 × 3 | Yes | DV | Yes | Uniformly dark, with grayish, hassle and black hues. | Pale/light bluish gray. | Yes |
G. festae | |||||||
100+ | 5–20 × 1.5–4 | Yes | V | Yes | Brown, grayish or yellow tones, with or without a clear midline; uniform granular pigmentation or distributed laterally. | Uniformly grayish-yellowish. | Yes |
G. rincona | |||||||
9 | 12 × 1.5 | Yes | V | ? | Grayish brown or grayish yellow, sometimes with a pale midline. | White. | No |
R. patagonicus | |||||||
65+ | 3–6 × 1–3.5 | Some | D | ? | Yellowish with black-brown in broad confluent spots/uniform dark brown-black. | White with darker blurry spots/uniform dark brown-black/pale brown. | No |
External appearance of freshwater planarians described from Chile. (A) Dugesia ambigua; (B) Dugesia similis; (C) Girardia canai; (D) Girardia chilla; (E) Girardia festae; (F) Girardia rincona; (G) Romankenkius patagonicus. Scale bars = 1 mm. Illustrations redrawn based on publications (
Regarding the copulatory apparatus, we summarized sketches based on reported information to highlight the differences between the species (Fig.
Reconstruction of the copulatory apparatus of species recorded for Chile. (A) G. canai; (B) G. chilla; (C) G. festae; (D) G. rincona; (E) R. patagonicus. Labels: bc = bursal canal, cb = copulatory bursa, cg = cement glands, ed = ejaculatory duct, go = gonopore, od = ovovitelline duct, pp = penis papilla, sd = sperm duct, se = false seminal vesicle, sg = shell glands, sv = seminal vesicle, vd = vas deferens. Sketches redrawn based on publications (
Synonyms. Planaria ambigua Böhmig, 1902; Curtisia ambigua
Distribution. Southern Chile (Magallanes Region).
The original description of the species was made by
The largest organism collected was about 7 mm long and 3 mm wide after fixation, but its copulatory organs were immature and the definitive configuration could not be concluded, thus the specimen was named Planaria ambigua. The mouth lies at the beginning of the posterior third and the genital gonopore is about 0.5 mm away from it. The ovaries are found in the anterior region and the testes lie dorsal between the third and fourth branches.
Synonyms. Planaria similis Böhmig, 1902.
Distribution. Central Chile (Valparaíso Region).
Very little information has been published on Dugesia similis. Many specimens were collected in 1893 in Salto brook (Fig.
The author commented that the animals are very similar in body conformation to the North American Planaria maculata Leidy, 1847 (a synonym of Girardia tigrina Girard, 1850): a slender anterior, acute-triangular head shape and pointed rear end (Fig.
Synonyms. none.
Distribution. Central Chile (Metropolitana and Valparaíso Regions).
The discovery of this species dates back to 1986. The authors’ particular interest was to find G. festae in the locations previously described by
The authors examined sixteen mature specimens and thus provided a very detailed description of the reproductive apparatus (Fig.
Synonyms. Dugesia chilla Marcus, 1954; Dugesia veneranda Martins, 1970.
Distribution. Central and South Chile (Valparaíso, Los Lagos and Magallanes Regions), Brazil, Argentina.
G. chilla has some peculiar characteristics, including a stubby cone or ball-shaped penis papilla and dorsal or dorsoventral testes. The first description of this species was made by
The specimens are slender, measuring approximately 9–25 mm long and 2–4.5 mm wide. The head is large, narrow, and pointy with protruding flexible auricles. Dorsal pigmentation is uniformly dark, dense, with grayish, hassle and black hues (Fig.
Marcus’s specimens from Pichi Pilluco Estuary, Llanquihue, were sexually mature, but the samples were contracted, and reconstruction of its copulatory apparatus was criticized by
The species seems to be distributed across South America. G. chilla has also been found in two locations in Southern Argentina: Moreno Lake at 765 masl and Ñireco River (
Synonyms. Planaria festae Borelli, 1898; Planaria dimorpha Böhmig, 1902; Planaria polyorchis Fuhrmann, 1914; Euplanaria aurita de Beauchamp, 1939; Dugesia titicana Hyman, 1939; Dugesia polyorchis Kenk, 1941; Dugesia dimorpha Marcus, 1954; Dugesia festai Marcus, 1954; Dugesia (Girardia) sanchezi Hyman, 1959; Dugesia (Girardia) polyorchis Ball, 1974; Girardia festai Sluys, 1992.
Distribution: Central and Southern Chile (Valparaíso, Metropolitana, Biobío and Los Ríos Regions), Argentina, Peru, Ecuador, Bolivia, Venezuela, Colombia, Curaçao, Brazil, Uruguay, Paraguay.
Girardia festae is one of the planarians present in Chile with the largest distribution and several synonyms. It has been found in springs, streams and lakes at altitudes up to 4800 masl. Its Latin American distribution has been summarized elsewhere (
The species reaches dimensions of 10–20 mm long and 1.5–4 mm wide. It has a triangular head with pointed auricles, typical of the genus Girardia. The body has the same width up to the rear third of the body, then narrows into a pointy posterior end. The dorsal pigmentation is variable, generally brown that varies towards grayish or yellow tones, with or without a clear longitudinal midline, plus a pattern of granular pigmentation that can be uniform throughout the body or distributed in the lateral regions (Fig.
The reproductive system of G. festae (Fig.
Synonyms. Dugesia rincona Marcus, 1954; D. (G.) rincona Ball, 1974; Girardia rincona
Distribution. Northern Chile (Coquimbo Region), Peru, Venezuela.
A detailed description of the reproductive apparatus (Fig.
The species was redescribed by Kawakatsu and coworkers based on material from Peru and Venezuela (
Synonyms. Planaria patagonica Borelli, 1901; Planaria michaelseni Böhmig, 1902; Curtisia patagonica Kenk, 1930; Curtisia michaelsoni Kenk, 1930; Cura patagonica Ball, 1974; Cura michaelseni Ball, 1974; Dugesia patagonica Ball, 1974.
Distribution. Southern Chile (Magallanes Region), Argentina.
The first description of Romankenkius patagonicus dates from the beginning of the 20th century.
Subsequently, expeditions by Haga in 1966 and Inoue in 1971 resulted in the collection of specimens of this species in seven locations in Torres del Paine, Tierra del Fuego (Fig.
Out of the sixty-five specimens collected in the description of
The dorsal surface of most of the specimens is covered by spots of a very dark brown or gray color, except for a pale colored median longitudinal line, which starts from the top of the eyes and extends almost to the posterior end of the body (Fig.
The specimens have a pair of small ovaries located ventrally between the second and third intestinal diverticula, while the testes are located mainly dorsally and pre-pharyngeally (
In 1954, Marcus wrote “Chile is one of the places whose Turbellaria fauna has become known much more through the collective exploitation of traveling zoologists, than through the activities of workers resident in the country itself”. Almost 70 years later, this statement remains valid. The information compiled for the species summarized in this paper, with records dating from the end of the 19th century and sparsely reaching the present, and mostly provided by non-Chilean scientists, reflects on the lack of specialists, interest, and/or resources in studying the freshwater planarians in this country. Furthermore, none of the reported specimens are deposited in Chilean museums. Herein, we provide a map with the record locations (Fig.
We have found some inconsistencies. The WoRMS database finds five matching records for Dugesiidae in Chile, but two of those actually correspond to one species (D. michaelsoni = R. patagonicus) and it lacks the records of D. ambigua, D. similis, and G. canai. We have revised some coordinates (Table
Chile represents a great source of biodiversity that should be carefully studied due to its different climates and biogeography; several hot spots of endemicity are found and must be protected. It could be estimated that there are many planarian species that have not yet been described for marine, terrestrial and freshwater environments, including cave planarians, which are surely present in Chile but have not been recorded so far. Records of Chilean land planarians were compiled by
In the future, we aim to welcome new investigators to resample type localities and carry out a thorough sampling to extend the knowledge of the species listed here. The correct determination of species in the family Dugesiidae is difficult due to similarities in both the external morphology and reproductive apparatus. Therefore, new studies should integrate the traditional techniques (morphological, anatomical and reproductive system description) with molecular analyses of nuclear and mitochondrial markers, such as ribosomal subunit 18 (18S), ribosomal subunit 28 (28S), cytochrome c oxidase subunit 1 (COI), elongation factor 1α (EF), and internal transcribed spacer 1 (ITS-1) sequencing (
We have summarized the characteristics and brief sampling history of the seven freshwater triclads known from Chile and hypothesize that there are many more to discover. Interestingly, Chile, Argentina and Brazil share at least two species: G. chilla and G. festae (
We are thankful to Francisco Brusa, Universidad Nacional de La Plata, for providing information and support. We thank the World Register of Marine Species (WoRMS) and the Turbellarian Taxonomic databases. We also thank the reviewers for their constructive comments. This work was funded by the Agencia Nacional de Investigación y Desarrollo de Chile (ANID)/FONDECYT Postdoctoral Grant number 3180580 to CVD. MLA and CVD received funding from ANID - MILENIO - ICN2021_044.