Research Article |
Corresponding author: Giovanni Mussini ( gm726@cam.ac.uk ) Academic editor: Alan Myers
© 2024 Giovanni Mussini, Nicole D. Stepan, Gersey Vargas.
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:
Mussini G, Stepan ND, Vargas G (2024) Two new species of Hyalella (Amphipoda, Dogielinotidae) from the Humid Chaco ecoregion of Paraguay. ZooKeys 1191: 105-127. https://doi.org/10.3897/zookeys.1191.113840
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The freshwater amphipod genus Hyalella Smith, 1874 is widely distributed in the Neotropics, with several biogeographically restricted species and a high cryptic diversity throughout South America. Tens of species of Hyalella have been documented from nearby Brazil and Argentina, but no systematic record of the genus exists for Paraguay. Here we describe two new species of Hyalella: H. mboitui sp. nov. and H. julia sp. nov. from the Ñeembucú wetlands of southwestern Paraguay. Hyalella mboitui sp. nov. and H. julia sp. nov. are characterised by a dorsally smooth body, pigmented eyes, uropod 1 endopod with a curved seta, the dorsal margin of uropod 3 ramus without setae, and uropod 3 peduncle longer than wide and with six setae apically. The two species are distinguished by their diagnostic mouthparts, with a median serrated edge on the lacinia mobilis in H. mboitui sp. nov. and two elongated lateral denticles with a serrated edge in H. julia sp. nov., and by the presence of a pronounced cup for the dactylus on gnathopod 2 in H. mboitui sp. nov. In addition, they show differences in the number of articles on antennae 1 and 2, in the relative length of the pereiopods, and in the numbers and types of setae on their gnathopods and uropods 1–3. Hyalella mboitui sp. nov. and H. julia sp. nov. represent the first taxonomically documented occurrence of Paraguayan freshwater amphipods. These new taxa attest to the largely unmapped species richness of freshwater invertebrates in the Humid Chaco of Paraguay. This potential biodiversity hotspot is currently under threat from land conversion, highlighting the need for more systematic studies and effective conservation of the local invertebrate biodiversity.
Amphipoda, conservation, Hyalella, new species description, Paraguay, taxonomy
Amphipods are a diverse clade of peracaridan crustaceans inhabiting both marine and freshwater environments, where they represent an ecologically and taxonomically significant component of the planktonic and benthic invertebrate fauna (
This knowledge gap has potentially broad-ranging repercussions for conservation and habitat management. Amphipods, including Hyalella, sustain key links in matter and energy transfers in freshwater ecosystems, where they act as ecologically abundant grazers and detritivores, provide hosts for diverse micro- and macro-parasites, and serve as important prey items for both vertebrate and invertebrate predators (
Here we describe two new species of Hyalella from the Ñeembucú wetlands, part of the Humid Chaco ecoregion of southwestern Paraguay (
Freshwater invertebrate specimens were sampled from September 8, 2021 to June 5, 2023, as part of an environmental impact assessment led by Fundación Para La Tierra (PLT) under contract from the Ministry of Public Works and Communications (Ministerio de Obras Públicas y Comunicaciones, MOPC). Five field sites in total were sampled in and near the city of Pilar, Ñeembucú Department (Paraguay): Yegros Paso (26°51'51"S, 58°16'11"W), San Lorenzo (26°52'35"S, 58°18'40"W), Costanera (26°50'52"S, 58°18'51"W), Ring Road (26°52'31"S, 58°14'59"W) and Laguna Gadea (26°50'9"S, 58°18'46"W). Samples were collected using a Seine net in 100 m transects, fragmented into 10 mini-transects of 10 m each. Upon completing the 100 m transect, investigators returned to the beginning, completing as many transects as allowed in a 2-hour period of continuous sampling. There were two 2-hour periods at each site (a total of 4 hours per site), between 7:00 and 9:00, and again between 15:00 and 17:00. This was repeated every three months for 2021–2023. The invertebrate specimens collected were all placed in jars with 70% ethanol and transferred to the PLT laboratory [Centro IDEAL (Investigation, Development, Environmental Education and Leadership), Pilar] for examination and identified to the lowest taxonomic level permitted by the available literature.
Measurements for the two new species were taken under an AmScope Trinocular Stereo Zoom Microscope 3.5×–90× magnification with a millimetric scale. Representative specimens (male paratypes and female allotypes) were dissected using a scalpel, pincers and fine needles, and mounted on permanent slides for storage and drawing under an OMAX 40×–2000× LED Microscope with built-in camera. Our description follows the setal terminology of
Type specimens are stored in the Scientific Collection of PLT (Colección Científica de PLT, CCPLT) at Centro IDEAL in Pilar, Ñeembucú Department (Paraguay).
Order Amphipoda Latreille, 1816
Family Dogielinotidae Gurjanova, 1953
Subfamily Hyalellinae Bulycheva, 1957
Genus Hyalella Smith, 1874
Holotype
, male (Fig.
Hyalella mboitui sp. nov., Department of Ñeembucú, Paraguay. Paratype, male A habitus B antenna 1 C antenna 2 D upper lip E left mandible, with detail of lacinia and incisor rotated anticlockwise (E1) and clockwise (E2) F right mandible, with detail of lacinia and incisor rotated anticlockwise (F1) and clockwise (F2) G lower lip H maxilla 1 I maxilla 2 J maxilliped. Scale bars: 1 mm (A); 0.5 mm (B, C); 0.2 mm (D–J); 0.1 mm (E1, E2, F1, F2).
Paraguay, Department of Ñeembucú, Pilar, Ring Road field locality (26°52'31"S, 58°14'59"W).
Flagella of antennae 1 and 2 with 13–14 and 16–17 articles, respectively. Left mandible incisor toothed, 5-denticulate; left lacinia mobilis multi-denticulate, with median serrated surface and two prominent elongated denticles laterally. Gnathopod 1 propodus subtriangular, without triangular space between propodus and dactylus, with papposerrate setae on disto-anterior corner. Gnathopod 2 propodus with papposerrate setae on disto-posterior margin, with palm with pronounced cup for dactylus. Pereopod 5 shorter than other pereopods. Uropod 1 endopod with a curved seta. Uropod 3 ramus dorsal margin without setae. Uropod 3 peduncle with two cuspidate setae and four simple setae apically. Uropod 3 peduncle longer than wide (rectangular).
Male (Figs
Antenna 1 about 2.2× shorter than body length, 1.4× shorter than antenna 2, 1.8× longer than peduncle of antenna 2; peduncle 1.1× longer than head length; flagellum with 13–14 articles, 1.5× longer than peduncle; aesthetascs occurring distally after article 4 (Fig.
Antenna 2 about 1.5× shorter than body length; peduncle 1.6× longer than head length; articles 1 to 3 with several simple setae on distal margin; flagellum 1.7× longer than peduncle, with 16–17 articles, with basal article elongated, with several simple setae on distal margins, and with four simple setae apically (Fig.
Basic amphipodan mandibles (sensu
Upper lip distal margin truncate; distal border covered by setules on ventral and dorsal faces (Fig.
Maxilla 1 inner plate slender, 1.8× shorter than outer plate, with two apical papposerrate setae and several setules laterally. Outer plate with nine serrate setae and several setules (Fig.
Maxilla 2 inner and outer plates subequal in length and width. Inner plate with one papposerrate seta and several simple and serrate setae apically, and several setules on inner face; outer plate with several simple setae on apex and margin, longest apically (Fig.
Maxilliped inner plate 2.0× longer than wide, apically truncated, with two cuspidate setae, several simple setae apically, and several setules on inner margin, comb-scales absent; outer plate approximately 1.3× longer than inner plate, apically rounded, with several apical and lateral simple setae, comb-scales absent; palp approximately 2.1× longer than inner plate, with four articles; article 1 1.1× longer than wide, inner margin with few simple setae; article 2 1.5× longer than wide, inner margin with several simple setae, outer margin with few simple setae; article 3 1.8× longer than wide, inner and outer margins with several long simple setae; article 4 unguiform, 3.0× shorter than third article, 1.6× longer than wide, inner margin with several long simple setae, with distal simple seta, with distal nail and comb-scales absent (Fig.
Gnathopod 1 subchelate; coxal plate 2.1× wider than long, with several simple setae on anterior and posterior margins; basis with one simple seta on anterior margin and one on disto-posterior corner; ischium with few simple setae on disto-posterior corner; merus with few simple setae on posterior margin; carpus 1.3× longer than wide, 1.1× longer and wider than propodus, posterior lobe produced and forming scoop-like structure, pectinate margin with several serrate setae, comb-scales and polygonal pattern; propodus 1.3× longer than wide, hammer-shaped, with several simple setae on anterior margin, with several papposerrate setae on disto-anterior corner; palm slope oblique, with several simple setae, margin convex, disto-posterior corner with long simple setae, and with a pronounced cup for dactylus; dactylus claw-like, congruent with palm, without comb-scales (Fig.
Gnathopod 2 subchelate; coxal plate 1.6× wider than long; basis with few simple setae on posterior margin and one serrate seta on disto-anterior margin; ischium and merus with few simple setae on posterior margin; carpus 1.8× wider than long, posterior lobe slim, produced between merus and propodus, margin pectinate with several serrate setae; propodus ovate, 1.4× longer than wide, comb-scales absent; palm subequal to posterior margin of propodus, slope oblique, margin convex, slightly irregular, with several simple setae and cuspidate setae with accessory seta; disto-posterior corner with two small cuspidate setae and several papposerrate setae, and with a pronounced cup for dactylus; dactylus claw-like, congruent with palm, without comb-scales (Fig.
Pereopods 3 to 7 simple. Pereopod 4 (Fig.
Pleopods not modified, biramous, elongated; peduncle 4.0× longer than wide, 1.7× mean size of rami, with coupling spines distally; both rami multi-annulated, longer than peduncle, with articles decreasing in size distally, with several plumose setae (Fig.
Uropod 1 1.5× longer than uropod 2; peduncle 1.1× longer than longest ramus, with four cuspidate setae; inner ramus 1.4× longer than outer ramus, 4.9× longer than wide, with three dorsal cuspidate setae, with one long curved seta and four cuspidate setae apically; outer ramus with three dorsal cuspidate setae and four cuspidate setae apically (Fig.
Uropod 2 1.5× shorter than uropod 1; peduncle rectangular, subequal in length to outer ramus and 1.2× shorter than inner ramus, 2.0× wider than outer ramus and 1.5× than inner ramus, with two cuspidate setae; inner ramus slightly longer than outer ramus, with four cuspidate setae dorsally and three cuspidate setae apically; outer ramus with three cuspidate setae dorsally and four cuspidate setae apically (Fig.
Uropod 3 (Fig.
Telson entire, 1.1× longer than wide, apically rounded, without setae laterally, with five apical cuspidate setae (Fig.
Coxal gills sac-like present on pereonites 3 to 6; sternal gills tubular and present on pereonites 3 to 7.
Female (Figs
Antennae similar in shape to male. Antenna 1 flagellum with 10–11 articles. Antenna 2 flagellum with 16–17 articles.
Gnathopod 1 (Fig.
Gnathopod 2 (Fig.
Telson approximately as long as wide, with more convex lateral margins than in male, and with five cuspidate setae apically (Fig.
Uropod 1 similar in size and shape to male uropod 1, except for absence of curved seta.
Freshwater, epigean.
Paraguay, Department of Ñeembucú, Pilar. Field localities of Ring Road (26°52'31"S, 58°14'59"W) and San Lorenzo (26°52'35"S, 58°18'40"W).
In reference to Mbói Tu’i, one of the seven legendary monsters of Guaraní mythology and protector of wetlands and aquatic life. The species is named in Guaraní in honour of it being an endemic Paraguayan species.
Holotype
, male (Fig.
Paraguay, Department of Ñeembucú, Pilar, Yegros Paso field locality (26°51'51"S, 58°16'11"W).
Flagella of antennae 1 and 2 with 10–11 and 13–14 articles, respectively. Left mandible incisor toothed, 4-denticulate; left lacinia mobilis 3-denticulate, with short median denticle and two prominent elongated denticles with serrated margin laterally. Gnathopod 2 propodus with palm lacking pronounced cup for dactylus, without papposerrate setae, with cuspidate setae with accessory setae on disto-posterior corner. Pereopod 5 slightly longer than other pereopods. Uropod 1 endopod with a curved seta. Uropod 3 ramus dorsal margin without setae. Uropod 3 peduncle with six simple setae apically. Uropod 3 peduncle longer than wide (rectangular).
Male (Figs
Hyalella julia sp. nov., Department of Ñeembucú, Paraguay. Paratype, male A habitus B antenna 1 C antenna 2 D upper lip E left mandible with detail of lacinia and incisor rotated anticlockwise (E1) and clockwise (E2) F right mandible, with detail of lacinia and incisor rotated anticlockwise (F1) and clockwise (F2) G lower lip H maxilla 1 I maxilla 2 J maxilliped. Scale bars: 1 mm (A); 0.5 mm (B, C); 0.2 mm (D–J); 0.1 mm (E1, E2, F1, F2).
Antenna 1 about 3.2× shorter than body length, 1.3× shorter than antenna 2, 2.2× longer than peduncle of antenna 2; peduncle not surpassing head length; flagellum with 10–11 articles, 2.1× longer than peduncle; aesthetascs occurring distally after article 4 (Fig.
Antenna 2 about half of body length; peduncle 1.1× longer than head; articles 1 to 3 with several simple setae on distal margin, article 3 with several simple setae on lateral margin; flagellum 2.4× longer than peduncle, with 13–14 articles, with basal article elongated; articles with several simple setae on distal margins; four simple setae apically (Fig.
Basic amphipodan mandibles (sensu
Upper lip distal margin rounded, covered by several setules on dorsal and ventral faces (Fig.
Maxilla 1 inner plate slender, 1.4× shorter than outer plate, with two apical papposerrate setae and several setules laterally; outer plate with nine serrate setae (Fig.
Maxilla 2 inner plate 1.1× longer than outer plate; inner plate with one papposerrate seta and several simple and serrate setae apically, with several setules laterally; outer plate with several simple setae, longest apically, with several setules laterally (Fig.
Maxilliped inner plate 1.7× longer than wide, apically truncated, with three apical cuspidate setae and several simple setae, without comb-scales; outer plate 1.3× longer than inner plate, apically rounded, with several apical and lateral simple setae; palp 2.3× longer than inner plate, 1.7× longer than outer plate, with four articles; article 1 1.3× longer than wide, with strongly concave distal margin; article 2 1.0× longer than wide, with inner, outer, and distal margins with several long simple setae; article 3 1.2× longer than wide, with inner and outer margins with several simple setae; article 4 unguiform, 1.5× longer than wide, 2.0× shorter than third article, with distal simple setae, with distal nail and comb-scales absent (Fig.
Gnathopod 1 subchelate; coxal plate 1.9× wider than long; basis with one simple seta on inner margin and one on disto-posterior corner, ischium with few simple setae on disto-posterior corner; merus with several simple setae on posterior margin; carpus 1.5× longer than wide, 1.2× longer and 1.2× wider than propodus, with several simple setae on disto-anterior corner, some with accessory seta, with few simple setae on inner margin, with posterior lobe folded to form scoop-like structure, with pectinate margin with comb-scales, several serrate setae and polygonal pattern; propodus 1.6× longer than wide, hammer-shaped, with simple seta with accessory seta on anterior margin, with inner margin with several simple setae, with several long simple setae on disto-anterior corner; palm slope transverse, margin slightly concave, with many simple setae, with disto-posterior corner with cuspidate seta with accessory seta; dactylus claw-like, congruent with palm, without comb-scales (Fig.
Gnathopod 2 subchelate; coxal plate 1.8× wider than long; basis with one simple seta on anterior margin and several simple setae on disto-posterior margin; merus with several simple setae on posterior margin; carpus 2.0× wider than long, with one simple seta on inner margin and two on disto-anterior, with posterior lobe slim produced between merus and propodus, with posterior margin pectinate, with several serrate setae and comb-scales; propodus ovate, 1.4× longer than wide, with two simple setae on anterior margin; palm subequal to posterior margin of propodus, slope oblique, margin convex, with several long and short simple setae with accessory setae; disto-posterior corner with two cuspidate setae with accessory setae; very shallow cup for dactylus; dactylus claw-like, congruent with palm, without comb-scales (Fig.
Pereopods 3 to 7 simple. Pereopods 3 and 4 (Fig.
Pleopods not modified, biramous, elongated; peduncle 4.0× longer than wide, 1.5× shorter than mean size of rami, with coupling spines distally; both rami multi-annulated, longer than peduncle; articles decreasing in size distally in both rami; both rami with several plumose setae (Fig.
Uropod 1 1.3× longer than uropod 2; peduncle 1.1× shorter than outer ramus and 1.2× shorter than inner ramus, with three cuspidate setae; inner ramus 1.1× longer than outer ramus, with four cuspidate setae dorsally, and one long curved seta and five cuspidate setae apically; outer ramus with four cuspidate setae dorsally and five cuspidate setae apically (Fig.
Uropod 2 1.3× shorter than uropod 1; peduncle rectangular, 1.0× shorter than outer ramus and 1.3× than inner ramus, 2.4× wider than outer ramus and 1.3× than inner ramus, with one cuspidate seta dorsally; inner ramus 1.3× longer than outer ramus, with three cuspidate setae dorsally and three cuspidate setae apically; outer ramus with four cuspidate setae dorsally and four cuspidate setae apically (Fig.
Uropod 3 (Fig.
Telson entire, 1.1× longer than wide, with convex margins, and rounded apically, without setae laterally, and with five cuspidate setae apically (Fig.
Coxal gills sac-like present on pereonites 3 to 6; sternal gills tubular and present on pereonites 3 to 7.
Female (Figs
Antennae similar in shape to male. Antenna 1 flagellum with 10–11 articles. Antenna 2 flagellum with 11–12 articles.
Gnathopod 1 (Fig.
Gnathopod 2 (Fig.
Telson subequal in length and width, with more convex lateral margins than in male, and with five cuspidate setae, one with accessory seta.
Uropod 1 similar in size and shape to male uropod 1, except for absence of curved seta.
Freshwater, epigean.
Paraguay, Department of Ñeembucú, Pilar. Field locality of Yegros Paso (26°51'51"S, 58°16'11"W).
In honour of the late Don Julio Rafael Contreras, for his seminal studies of Paraguayan biodiversity and generous support of Fundación Para La Tierra.
Hyalella mboitui sp. nov. and H. julia sp. nov. can be recognised as distinct species based on the taxonomic keys by
Like H. brasiliensis Bousfield, 1996 from Paraná State (Brazil), both H. mboitui and H. julia lack plumose setae on their telson, but can be readily distinguished from this species by the number of setae on uropods 1 and 2 (
The two new Paraguayan taxa are also readily distinguishable from recently described Hyalella species from nearby Argentina (
Despite the geographical vicinity of their type locations, H. mboitui and H. julia are separated by clear morphological differences at the level of the gnathopods, uropods and mandibles, as well as by minor differences in the morphology and setal covers of their antennae, maxillae and maxillipeds. Antennae 1 and 2 have fewer articles in their flagellum in H. julia than in H. mboitui: H. julia has minimally 13 articles in antenna 1 and 16 in antenna 2, whereas H. mboitui has minimally 10 in antenna 1 and 13 in antenna 2. The mandibles of the two taxa differ in the number of incisor teeth, with 5 and 6 teeth present in the left mandibles of H. mboitui and H. julia, respectively, and 4 and 7 in their right mandibles. In addition, the left lacinia mobilis of H. julia lacks the distinctive median serrated surface of H. mboitui, and sports instead two prominent, elongated denticles with a serrated edge laterally. The maxillipeds of the two species differ in the number of cuspidate setae on the outer plate and in the shape of palp articles (Figs
The geographical vicinity of the two new species and their distinct mandibular morphologies suggest that their differences may stem at least in part from trophic partitioning (
The type locality of H. julia is managed for ongoing conservation and research projects on the endangered Pilar tuco-tuco (Ctenomys pilarensis). In contrast, major developments are scheduled or currently taking place at the type localities of H. mboitui, San Lorenzo and Ring Road, for the planned construction of flood defences. The connections between the bodies of water inhabited by H. mboitui and H. julia, and their seasonal continuity with the Ñeembucú River, preliminarily suggest that their area extends beyond the type localities. However, Hyalella is known for its high degree of endemism across South America, and the geographic range of different species in the genus is highly variable (
The discovery of two new crustacean species, collected in a non-targeted impact assessment survey near an urban area, highlights the untapped potential of the Ñeembucú wetlands for biodiversity and conservation research. This ecologically important patchwork of rivers, streams, and flooded grasslands is severely understudied, and its invertebrate fauna remains virtually unexplored amid escalating anthropic impacts (
We thank all the PLT interns, volunteers and staff members involved for their assistance throughout the project.
The authors have declared that no competing interests exist.
No ethical statement was reported.
GM acknowledges support from a NERC (Natural Environment Research Council) C-CLEAR DTP studentship [RG96579].
G.M. made specimen photographs and drawings and wrote the first draft of the manuscript. G.V. and N.D.S. provided field site photographs and assisted with manuscript drafting, editing, and specimens storage and cataloguing.
Giovanni Mussini https://orcid.org/0009-0002-1093-5865
Gersey Vargas https://orcid.org/0000-0002-5625-8284
All of the data that support the findings of this study are available in the main text.