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Checklist of species of Stratiodrilus Haswell, 1900 (Annelida: Histriobdellidae), and new host records from Southern Brazil
Species of Stratiodrilus is ectocommensal of freshwater decapod crustaceans, and it is also the most specious genus within the family Histriobdellidae. However, a global review of the species of Stratiodrilus and their hosts has not been performed, since the study of Steiner & Amaral (1999). Therefore, based on literature and on our data, we present a list of Stratiodrilus species and their hosts. We searched for host and occurrence records on papers of species descriptions, review papers and specialized databases. Forty-one host records were summarized worldwide. All host taxa were freshwater decapod crustaceans belonging to Aegla, Astacoides, Astacopsis, Cherax, Parastacus, Samastacus and Trichodactylus. In addition, we reportfour new host records for Stratiodrilus circensis, being Aegla lata the Northernmost host record for this polychaete species. Neotropical region harbors the greatest species richness of Stratiodrilus, although this genus is also found in Australia and Madagascar associated with crayfishes. Aegla was the most reported host genus associated with Stratiodrilus spp., as well as S. circensis was the most cited for different hosts. Thus, our aim in this paper is to contribute for an update of the known species of Stratiodrilus and also to discuss the distribution of these histriobdellids.
Key words: Stratiodrilus, decapod crustaceans, ectocommensal, checklist.
At least 292 species of commensal polychaetes belonging to 28 families are known. One of these belongs to Histriobdellidae Vaillant, a highly modified family of entirely symbiotic small polychaetes (Martin & Britayev 1998). Histriobdellidae is composed of three genera: Histriobdella van Beneden found on the eggs of squat lobsters from Europe and North America; Stratiodrilus Haswell, 1900, found inside the branchial chambers, internal part of abdomen, on the pleopods and egg masses of freshwater crabs and crayfishes from Australia, Madagascar and South America; and Steineridrilus Zhang, 2014 (=Dayus Steiner & Amaral, 1999), found on pleopods of marine isopods from South Africa (Steiner & Amaral 1999). Both Histriobdella and Steineridrilus are monospecific genera, but the first is found in a variety of hosts from Northern hemisphere (Brattey & Campbell 1985) while the second one is found in a single isopod host, Cirolana venusticauda var. simplex Barnard (Führ 1971). On the other hand, Stratiodrilus is the most specious genus, occurring in a variety of hosts and localities of Southern hemisphere (see Steiner & Amaral 1999).
Steiner & Amaral (1999) carried out the last revision of the family Histriobdellidae and summarized seven species of Stratiodrilus associated to decapod crustaceans of seven genera (Aegla Leach; Trichodactylus Latreille; Astacoides Guérin-Méneville; Cherax Erichson; Astacopsis Huxley; Parastacus Huxley; and Samastacus Riek). Furthermore, these authors described two new species: Stratiodrilus robustus Steiner & Amaral, 1999, associated with Trichodactylus sp. from São Paulo State, Southeastern Brazil; and Stratiodrilus circensis Steiner & Amaral, 1999, associated with Aegla sp. from Paraná State, Southern Brazil.
The last described species (Stratiodrilus brevicirrus Amato, Daudt & Amato, 2004, associated with Aegla platensis Schmitt, 1942, from Rio Grande do Sul State, Southern Brazil) as well as the key for eleven species of Stratiodrilus knownto date were proposed by Amato et al. (2004). Although Stratiodrilus is the most reported genus in the literature, there are no recent studies on global diversity of this ectocommensal and their hosts.
Thus, we provide a checklist of species of Stratiodrilus and their hosts, as well as the distribution range of S. circensis with four new host records and its ecological descriptors.
Material and methods
Bibliographic search. The checklist of species of Stratiodrilus was based on original papers of species descriptions, on all periodical journals and/or books for this taxon, mainly on Martin and Britayev (1998), Steiner and Amaral (1999), and Amato et al. (2004). The bibliographic research was carried out up to November, 2017. The following databases were used in order to ensure that we have compiled all available information: ISI Web of Knowledge, Google Scholar, World Register of Marine Species, Information System on Brazilian Biodiversity (SiBBr), Global Biodiversity Information Facility (GBIF), and Integrated Taxonomic Information System.
For each record we present the species of Stratiodrilus, authority name, and year. We also inform the country where the species of Stratiodrilus were found and, when possible, the locality records for each host species associated. Records of Stratiodrilus spp.and hosts associated were arranged in alphabetical order. Records whose epithets were not informed by the authors were lelft for last. We present the geographic coordinates for each locality when they were informed in the original record; when they were not informed, we assigned geographic coordinates using Google Earth. Abbreviations for localities cited in this paper are:
Argentina (ARG): Província de Tucumán (TUC).
Australia: New South Wales (NSW); Queensland (QLD); Tasmania (TAS).
Brazil: Paraná State (PR); Rio Grande do Sul State (RS); São Paulo State (SP).
Chile (CHI): Central Chile (CNC); Concepción (CON).
Uruguay: Departamento de Canelones (CNL).
The map of the geographic distribution of S. circensis was built with ArcMap 10.5.1, application of the ESRI ArcGIS program package (student testing license: EVA465642350), using Geodetic Datum WGS84.
Field work. Collects of aeglid hosts were made in four streams belonging to three river basins of the Paraná State. Aegla castro Schmitt, 1942 was monthly collected from June 2012 to May 2013 in the Couro stream (23º57’15”S, 51º07’00”W), Tibagi river basin; Aegla lata Bond-Buckup & Buckup, 1994was monthly collected from August 2014 to June 2015 in the Bule stream (23º26’60”S, 51º15’06”W), a tributary of the Apertados river, Tibagi river basin; Aegla parana Schmitt, 1942 was collected in June 2017 in the Pinhão river (25º33’14”S, 51º26’23”W), Iguaçu river basin; and, Aegla sp. was monthly collected from July 2015 to October 2015 in the Lageadão river (24º01’33”S, 51º36’47”W), Ivaí river basin.
Hosts sampling was made using hand nets of 90 cm diameter and 1 mm mesh size. Rocks and leaf litter of the bottom substrate were removed while another researcher placed a hand net downstream to collect the crustaceans. All hosts were immediately individualized in bottles containing stream water.
In the laboratory, to remove and prevent the body contraction of polychaetes, the hosts were immersed in stream water with menthol (Amato et al. 2004), and then, they were analized under stereomicroscopy to actively search, and had their specimens of Stratiodrilus removed from the exoskeleton and internal part of abdomen. In addition, stream water was checked by filtrating using a 250 m mesh size in order to collect symbionts detached from the body of hosts. Specimens of Stratiodrilus were counted and some were fixed in AFA (solution of ethanol 70%, formaldehyde, and acetic acid), following procedures described by Amato et al. (1991).
Ecological descriptors such as prevalence (P%), mean abundance (MA), and mean intensity (MI) were calculated according to Bush et al. (1997) for each species of host.
The specimens analyzed were deposited in the Collection of Polychaetes of the Museu de Zoologia “Adão José Cardoso”, of the Instituto de Biologia of the Universidade Estadual de Campinas (ZUEC POL), São Paulo State, Brazil: lot 1, 2 male specimens, ZUEC POL 21313; lot 2, 2 female specimens, ZUEC POL 21314; lot 3, 3 male specimens, ZUEC POL 21315; lot 4, 3 female specimens, ZUEC POL 21316; lot 5, 2 male specimens, ZUEC POL 21317; lot 6, 2 female specimens, ZUEC POL 21318; lot 7, 1 male specimen, ZUEC POL 21319; and lot 8, 1 female specimen, ZUEC POL 21320.
Literature review and our own data allowed us to compile 11 species of Stratiodrilus worldwide associated with 41 host records (Table 1), all of which being freshwater decapod crustaceans. In Madagascar, there is an endemic species recorded being associated with only one host species. Two species of Stratiodrilus are endemic to Australia, being associated with five different host taxa. On the other hand, eight species of Stratiodrilus are endemic to Neotropics, and 35 records of hosts are reported for this region. In Argentina, Brazil, Chile and Uruguay were found, respectively, two, six, four, and one species of Stratiodrilus.
We report A. castro, A. lata, A. parana and Aegla sp. as new host records for S. circensis. Furthermore, we extend the distribution geographic range for S. circensis and we report the northernmost occurrence record for this polychaete species (23º26’59.80”S, 51º15’5.81”W) (Figure 1), as well as we refer Tibagi and Ivaí river basins as new drainages for its occurrence.
A total of 586 specimens of A. castro were collected (285 females and 301 males), of which 53.92% had specimens of S. circensis associated, 2,326 specimens on female crabs and 2,249 specimens on male crabs. Mean intensity was 14.48±1.07 specimens of S. circensis, whereas the mean abundance was 7.81±0.65. Seventy-four specimens of A. lata were collected (41 females and 33 males), of which 62.16% had specimens of S. circensis, 133 specimens on female crabs, and 119 specimens on male crabs. Mean intensity was 5.48±0.77 specimens of S. circensis, whereas the mean abundance was 3.40±0.57. Ten specimens of A. parana were collected (3 females and 7 males), of which 80.00% had specimens of S. circensis, 12 specimens on female crabs, and 32 specimens on male crabs. Mean intensity was 5.50±1.61 specimens of S. circensis, whereas the mean abundance was 4.40±1.47. Finally, 24 specimens of Aegla sp. were collected (8 females and 16 males), of which 95.83% had specimens of S. circensis, 175 specimens on female crabs, and 322 specimens on male crabs. Approximately, 21.61±5.55 specimens of S. circensis were associated with each host, whereas the mean abundance was 20.71±5.39.
Subsequently, the species of Stratiodrilus and their hosts, as well as new records are listed as the following.
Family Histriobdellidae Vaillant
Stratiodrilus Haswell, 1900
Stratiodrilus aeglaphilus Vila & Bahamonde, 1985
Type-locality: Maípo River, Talagante area, Central Chile, Chile.
Host and locality records: Aegla laevis (Latreille)—Maípo River, Talagante area (CNC) (Vila & Bahamonde 1985); Aguas Claras stream (CNC) (Vila & Bahamonde 1985; Pardo et al. 2008). Aegla laevis laevis (Latreille)—Peñaflor tributary, Maípo River (CNC) (Vila & Bahamonde 1985). Aegla laevis talcahuano Schmitt, 1942—“Estero” Bellavista between Lirquén and Tomé (CON) (Moyano et al. 1993). Unspecified species of Aeglidae and Parastacidae—from Petorca to Chiloé (CHI) (Steiner & Amaral 1999).
Stratiodrilus arreliai Amaral & Morgado, 1997
Type-locality: Peroba Cave, Municipality of São Pedro, State of São Paulo, Brazil.
Host and locality records: Aegla perobae Hebling & Rodrigues—Peroba Cave, municipality of São Pedro (SP) (Amaral & Morgado 1997). Aegla sp.—Creeks near Jaraguá Peak, municipality of São Paulo (SP) (Amaral & Morgado 1997).
Stratiodrilus brevicirrus Amato, Daudt & Amato, 2004
Type-locality: Mineiro creek, locality of Fazenda Fialho, Municipality of Taquara, State of Rio Grande do Sul, Brazil.
Host and locality records: Aegla platensis Schmitt, 1942—Mineiro creek, Taquara (RS) (Amato et al. 2004).
Stratiodrilus circensis Steiner & Amaral, 1999
Type-locality: Piraquara River, Municipality of Piraquara, State of Paraná, Brazil.
Host and locality records: Aegla abtao Schmitt, 1942—Toltén River (CHI) (Steiner & Amaral 1999). Aegla bahamondei Jara—Toltén River (CHI) (Steiner & Amaral 1999). Aegla castro Schmitt, 1942—Couro stream, Tibagi river basin (PR) (Present study). Aegla laevis (Latreille)—Riñihue Lake (CHI) (Steiner & Amaral 1999); Negro river and lakes in Parque Nacional de Nahuel Huapí (ARG) (Steiner & Amaral 1999). Aegla lata Bond-Buckup & Buckup, 1994—Bule stream, a tributary of the Apertados river, Tibagi river basin (PR) (Present study). Aegla leptodactyla Buckup & Rossi—Divisa river, a tributary of the Pelotas river, municipality of São José dos Ausentes (RS) (Daudt & Amato 2007). Aegla neuquensis neuquensis (Schmitt, 1942)—Chico river (TUC) (Steiner & Amaral 1999). Aegla parana Schmitt, 1942—Pinhão river, Iguaçu river basin (PR) (Present study). Aegla schmitti Hobbs III—Carvalho river, a tributary of Piraquara river, Parque Nacional Guaricana (PR) (Daudt & Amato 2007). Aegla sp.—Riñihue Lake (CHI) (Steiner & Amaral 1999). Aegla sp.—Piraquara river (PR) (Steiner & Amaral 1999). Aegla sp.—Lageadão river, Ivaí river basin (PR) (Present study). Unspecified species of Aeglidae and Parastacidae—from Petorca to Chiloé (CHI) (Steiner & Amaral 1999).
Distribution: Argentina, Brazil, Chile.
Stratiodrilus haswelli Harrison, 1928
Type-locality: Not informed in the original description.
Host and locality records: Astacoides madagascariensis (Milne Edwards & Audouin) (Uninformed locality) (Harrison 1928).
Stratiodrilus novaehollandiae Haswell, 1913
Type-locality: Probably Blue Mountains, New South Wales, Australia.
Host and locality records: Astacopsis serratus (Shaw)—Blue Mountains (NSW); Murrumbidgee River, Murray River System (NSW) (Haswell 1913). Cherax dispar Riek—Gap Creek, Mount Coottha (QLD) (Cannon & Jennings 1987). Cherax punctatus Clark—Gap Creek, Mount Coottha (QLD) (Cannon & Jennings 1987).
Stratiodrilus platensis Cordero, 1927
Type-locality: Probably Solís Chico and Miguelete streams, Departamento de Canelones, Uruguay.
Host and locality records: Aegla laevis (Latreille)—Solís Chico and Miguelete streams (CNL) (Cordero 1927). Samastacus spinifrons (Philippi)—Chile (Uninformed locality) (Rudolph 2002). Aegla sp.—Lakes of Parque Nacional Nahuel Huapí (ARG) (Dioni 1972). Parastacus sp.—Lakes of Parque Nacional Nahuel Huapí (ARG) (Dioni 1972). Trichodactylus sp.—Creeks near Cananéia (SP), Southeast Brazil (Steiner & Amaral 1999).
Distribution: Argentina, Brazil, Chile and Uruguay.
Stratiodrilus pugnaxi Vila & Bahamonde, 1985
Type-locality: Reumén, Valdivia, Chile.
Host and locality records: Parastacus pugnax (Poeppig)—Reumén (VAL) (Vila & Bahamonde 1985). Parastacus sp.—Andalién River, Chaimávida (CON) (Moyano et al. 1993). Unspecified species of Aeglidae and Parastacidae—from Petorca to Chiloé (CHI) (Steiner & Amaral 1999).
Stratiodrilus robustus Steiner & Amaral, 1999
Type-locality: Ribeira Valley, State of São Paulo, Brazil.
Host and locality records: Trichodactylus sp.—Streams near Ribeira Valley (SP) (Steiner & Amaral 1999).
Stratiodrilus tasmanicus Haswell, 1900
Type-locality: Hobart, Tasmania, Australia.
Host and locality records: Astacopsis franklinii (Grey)—Hobart (TAS) (Haswell 1900). Astacopsis franklinii var. tasmanicus Erickson—Hobart (TAS) (Haswell 1900).
Stratiodrilus vilae Amato, 2001
Type-locality: Municipality of Taquara, State of Rio Grande do Sul, Brazil.
Host and locality records: Parastacus brasiliensis (von Martens)—Tributary of the Mineiro creek, Taquara (RS) (Amato 2001); unidentified creek of Mariana Pimentel (RS) (Amato 2001). Parastacus defossus Faxon—Swampy area in Mariana Pimentel (RS) (Amato 2001).
Histriobdellids are small polychaetes that stablish obligatory symbiotic interactions with different groups of hosts (Martin & Britayev 1998). Unlike Histriobdella and Steineridrilus,which are both monospecific genera and exclusively associated with marine hosts, all species of Stratiodrilus are associated only with freshwater decapod crustaceans (Steiner & Amaral 1999). Furthermore, Stratiodrilus is undoubtedly the genus with the highest species richness, as well as the one with the highest number of occurrence records and different hosts associated.
Anomuran freshwater crabs of the genus Aegla are the most reported host species to be associated with Stratiodrilus (21 host records), followed by Parastacus (5), Astacopsis (3), Cherax (2), Trichodactylus (2), Astacoides (1), and Samastacus (1). In addition, the literature review has reported other records whose species of Aeglidae (3 records) and Parastacidae (3 records) were not informed. The most reported aeglid species to establish association with Stratiodrilus is Aegla laevis (6 records), although some of these records are misleading identifications of hosts, such as studies that reported A. laevis to the East of the Andes.
In this paper, we presented the second occurrence record of S. circensis in the State of Paraná. We report four new hosts, being A. lata associated with S. circensis the northernmost host record reported in literature. Moreover, we presented ecological descriptors of all species collected, being Aegla sp. from Lageadão river the host species with the highest numbers of prevalence, mean abundance, and mean intensity of infestation. In contrast, A. castro presented the lowest prevalence, while A. lata and A. parana presented the lowest mean abundance and the lowest mean intensity of infestation, respectively. However, there are few studies on Stratiodrilus spp. that address ecological descriptors, thus that would be interesting to investigate probable differences in the abundance of this polychaetes among several host populations.
The conspicuous distribution of Stratiodrilus spp.suggests the idea of a gondwanan origin and a single colonization of freshwaters by this group (Harrison 1928) during Palaeozoic or Mesozoic—as well as the polychaete Namanereis Chamberlin, that also succeeded in invading freshwaters at that period (Glasby & Timm 2008). Hence, as these ectocommensals are found exclusively in freshwaters from separated land masses, Harrison (1928) suggested that Stratiodrilus, or its ancestral, have colonized inland waters associated with ancestral parastacid crayfishes that have migrated to the continent before the breakup of the Gondwana. After many decades, molecular studies revealed that parastacid crayfishes constitute a monophyletic group and strongly supported that their origin occurred only once in Pangea during the Triassic (185–225 million years ago) (Crandal et al. 2000), corroborating the Harrison’s hypothesis. Moreover, it is interesting to note that among the hosts that are reported to establish association with Stratiodrilus (families Aeglidae, Parastacidae and Trichodactylidae), only parastacid crayfishes are found simultaneously in the three large biogeographic regions (Neotropical, Afrotropical, and Australasian).
Nevertheless, once the continents were separated, Stratiodrilus may have cospeciated together with their parastacid hosts, since the composition ectocommensal-host associations of each biogeographic region is unique. Cospeciation events have been reported in the literature for many host-symbiont systems (lice-rodent host: Hafner et al. (1994); oxyurid-primate host: Hugot (1999); myzostomid annelids-crinoid host: Lanterbecq et al. (2010); digenean-fish host: Martínez-Aquino et al. (2013; 2014)), supporting a congruence between parasite and host evolutionary and biogeographic histories. This pattern was also demonstrated for temnocephalids (Martínez-Aquino et al. 2016), another group of ectocommensals of gondwanan origin associated with freshwater decapods. Likewise, as the diversity of Stratiodrilus may be associated to the distribution of their hosts (Amaral & Morgado 1997), some ectocommensal-host associations that are unique from each biogeographic region are presented in our checklist. For example, S. arreliai is exclusively found in association with Astacoides spp., endemic crayfishes from Madagascar (Crandal et al. 2000; Jones et al. 2007); S. tasmanicus is found only in association with Astacopsis spp., endemic crayfishes from Australia (Crandal et al. 2000; Tadashi & Crandall 2016); and several species of Stratiodrilus—of which do not occur neither in Australia or Madagascar—are found in association with Parastacus spp., endemic crayfishes from South America (Bond-Buckup 2003). Besides, in order to test the possible effects of host speciation on ectocommensal speciation, studies based on biogeographic methods and molecular data of histriobdellids are strongly encouraged.
Stratiodrilus is widely distributed in Southern hemisphere although the highest species richness is in the Neotropics. It is also interesting to note that the most reported host genus (21 records) which occurs in association with Stratiodrilus is Aegla. These records may be due either to the lack of studies on histriobdellids in Australia and Madagascar, or probably an indication that the speciation of Stratiodrilus spp. occurred more intensely in the Neotropics after aeglids radiation, when several marine transgressions favored the dispersion and speciation of these populations during the early formation of the Andes Cordillera (Pérez-Losada et al. 2004). Thus, the high richness of Stratiodrilus spp., coupled with their association with aeglids—the only anomuran taxon entirely adapted to freshwater habitats (Bond-Buckup et al. 2008)—may suggest the existence of coevolutionary events between ectocommensals and aeglid hosts.
Despite of most species of Stratiodrilus are found in a single country, S. platensis isfound in four countries, five localities, one of them unspecified (S. platensis in association with S. spinifrons from Chile). Stratiodrilus platensis was previously reported to occur in association withA. abtao, A. bahamondei, and A. neuquensis neuquensisby Lang (1949), Roubaud (1962), Vila & Bahamonde (1985), Moyano et al. (1993) and Amaral & Morgado (1997). However, these host records are probably uncertain on account of the studies of Steiner & Amaral (1999) and Daudt & Amato (2007) which S. platensis was considered as S. circensis. Likewise, A. laevis as host of S. circensis was incorrectly reported from Uruguay (Cordero 1927) and Argentina (Lang 1949), since Bond-Buckup (2003) stablished that this speciesdoes not occur in the Eastern Andes. Yet, other authors suggested that these specimens should be probably A. platensis, A. uruguayana,or A. prado (Amato 2001; Amato et al. 2004). Furthermore, Aegla sp. from Piraquara river (PR) associated with S. circensis was suggested to be A. schmitti by Amato (2001).
Some species of Stratiodrilus were also misidentified, which, however, were corrected by later studies. For instance, Steiner & Amaral (1999) and Daudt & Amato (2007) referred S. circensis instead of S. platensis associated with A. neuquensis neuquensis from Argentina and A. bahamondei, A. abtao and A. laevis from Chile. Therefore, S. circensis is currently the species of the genus most cited for different hosts (14) as well as the most recorded to different geographic localities (14), being found along Argentina, Brazil and Chile (Daudt & Amato 2007).
In any case, the records of S. circensis for Argentina and Chile should be carefully considered, since some authors neither deposit the studied specimens in scientific collections nor mentioned that they studied deposited species (Daudt & Amato 2007). Furthermore, according to the literature review the distribution of S. circensis is wide and extends to hydrographic basins of different geological ages. For this reason, more studies using molecular tools are needed to verify whether S. circensis can be a complex of cryptic species.
Here, we contribute to an information update of known species of Stratiodrilus, as well as to the increase of histriobdellid-host associations reports, which may represent a source of information on life history of this polychaete and their hosts However, because Stratiodrilus is a group scarcely studied, we encourage molecular and phylogeographic approaches on this genus within Histriobdellidae to infer on biogeographic and coevolutionary patterns of these organisms with their hosts.
We would like to thank colleagues from LabIAS (Laboratório de Invertebrados Aquáticos e Simbiontes) for the help in fieldwork, mainly to researcher Drª. Cecilia M. Guerrero-Ocampo; to Ana Carolina Vizintim for the help with the map; and to Areta E. B. Cordeiro for the help with English review. We are thankful to MZUEL (Museu de Zoologia da Universidade Estadual de Londrina) for making available vehicles for samplings; to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the scholarship given to the authors.
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FIGURE 1. Geographical distribution range of species of Stratiodrilus worldwide. Numbers inside the countries indicate the number of species recorded. Black circles refer to localities where Stratiodrilus circensis Steiner & Amaral, 1999 has been found. Geodetic datum WGS84.
TABLE 1. Species of Stratiodrilus Haswell, 1900 and hosts reported in the world. UN indicates uninformed locality.
|Biogeographic region||Country||Species of Stratiodrilus||Hosts||Geographic coordinates||References|
|Afrotropical||Madagascar||Stratiodrilus haswelli||Astacoides madagascariensis||UN||Harrison (1928)|
|Australasian||Australia||Stratiodrilus tasmanicus||Astacopsis franklinii||*42º52’S, 147º19’E||Haswell (1900)|
|Astacopsis franklinii var. tasmanicus||*42º52’S, 147º19’E||Haswell (1900)|
|Stratiodrilus novaehollandiae||Astacopsis serratus||*33º29’S, 150º14’E||Haswell (1913)|
|Cherax dispar||*27º27’S, 152º56’E||Cannon & Jennings (1987)|
|Cherax punctatus||*27º27’S, 152º56’E||Cannon & Jennings (1987)|
|Argentina||Stratiodrilus platensis||Aegla sp.||*40º54’S, 71º30’W||Dioni (1972)|
|Parastacus sp.||*40º54’S, 71º30’W||Dioni (1972)|
|Stratiodrilus circensis||Aegla laevis1||*40º54’S, 71º30’W||Steiner & Amaral (1999)|
|Aegla neuquensis neuquensis||*27º17’S, 65º15’W||Steiner & Amaral (1999)|
|Brazil||Stratiodrilus platensis||Trichodactylus sp.||*25º02’S, 47º58’W||Steiner & Amaral (1999)|
|Stratiodrilus arreliai||Aegla perobae||22º55’00”S, 47º87’00”W||Amaral & Morgado (1997)|
|Aegla sp.||*23º27’S, 46º45’W||Amaral & Morgado (1997)|
|Stratiodrilus circensis||Aegla castro #||23º57’15”S, 51º07’00”W||Present paper|
|Aegla lata #||23º26’60”S, 51º15’06”W||Present paper|
|Aegla leptodactyla||28º38’17”S, 49º57’46”W||Daudt & Amato (2007)|
|Aegla parana #||25º33’14”S, 51º26’23”W||Present paper|
|Aegla schmitti||25º46’23”S, 48º51’25”W||Daudt & Amato (2007)|
|Aegla sp.||*25º28’27”S, 49º06’34”W||Steiner & Amaral (1999)|
|Aegla sp. #||24º01’33”S, 51º36’47”W||Present paper|
|Stratiodrilus robustus||Trichodactylus sp.||*24º30’43”S, 47º29’13”W||Steiner & Amaral (1999)|
|Stratiodrilus vilae||Parastacus brasiliensis||29º37’S, 50º47’W; 30º21’S, 51º35’W||Amato (2001)|
|Parastacus defossus||*30º21’S, 51º35’W||Amato (2001)|
|Stratiodrilus brevicirrus||Aegla platensis||29º46’00”S, 50º53’00”W||Amato et al. (2004)|
|Chile||Stratiodrilus platensis||Samastacus spinifrons||UN||Rudolph (2002)|
|Stratiodrilus aeglaphilus||Aegla laevis||*33º39’54”S, 70º55’42”W; 33º44’27”S, 70º53’60”W||Vila & Bahamonde (1985); Pardo et al. (2008)|
|Aegla laevis laevis||33º45’00”S, 70º45’00”W||Vila & Bahamonde (1985)|
|Aegla laevis talcahuano||*36º40’00”S, 72º56’40”W||Moyano et al. (1993)|
|Aeglidae gen. sp.||UN||Steiner & Amaral (1999)|
|Parastacidae gen. sp.||UN||Steiner & Amaral (1999)|
|Stratiodrilus pugnaxi||Parastacus pugnax||39º45’S, 73º45’W||Vila & Bahamonde (1985)|
|Parastacus sp.||*36º47’00”S, 72º56’04”W||Moyano et al. (1993)|
|Aeglidae gen. sp.||UN||Steiner & Amaral (1999)|
|Parastacidae gen. sp.||UN||Steiner & Amaral (1999)|
|Stratiodrilus circensis||Aegla abtao||*39º15’S, 72º16’W||Steiner & Amaral (1999)|
|Aegla bahamondei||*39º15’S, 72º16’W||Steiner & Amaral (1999)|
|Aegla laevis||*39º48’03”S, 72º21’60”W||Steiner & Amaral (1999)|
|Aegla sp.||*39º48’03”S, 72º21’60”W||Steiner & Amaral (1999)|
|Aeglidae gen. sp.||UN||Steiner & Amaral (1999)|
|Parastacidae gen. sp.||UN||Steiner & Amaral (1999)|
|Uruguay||Stratiodrilus platensis||Aegla laevis2||*34º38’14”S, 55º43’48”W; *34º48’36”S, 56º11’53”W||Cordero (1927)|
1 A. laevis is not found to the east of the Andes according to Bond-Buckup (2003).
2 A. laevis is not found to the east of the Andes according to Bond-Buckup (2003). Species suggested to be A. platensis, A. prado or A. uruguayana (Amato 2001; Amato et al. 2004).
* coordinates not informed in the original paper, but obtained for this study using Google Earth.
# New host record
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