Research Article |
Corresponding author: Juliana Lopes Segadilha ( julianasegadilha@gmail.com ) Academic editor: Sammy De Grave
© 2020 Juliana Lopes Segadilha, Cristiana Silveira Serejo.
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:
Segadilha JL, Serejo CS (2020) First records of Pseudozeuxidae and Metapseudinae (Metapseudidae) (Crustacea, Tanaidacea) in Southwestern Atlantic, with descriptions of two new species. Zoosystematics and Evolution 96(2): 723-745. https://doi.org/10.3897/zse.96.56097
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Based on specimens collected from eulittoral zone in rocky shores of northeast of Rio de Janeiro (Brazil) on 2017, two new tanaidaceans species from two different suborders are described: Apseudomorpha brasiliensis sp. nov. (Apseudomorpha, Metapseudidae) and Pseudozeuxo fischeri sp. nov. (Tanaidomorpha, Pseudozeuxidae). Diagnostic characters of Apseudomorpha brasiliensis are mandible palp article-2 and article-3 with six and nine finely penicillate setae on inner margin, respectively; pereopod-1 carpus and propodus with two and four ventral spines, respectively; pleonites 2 and 5 with pleura having long distal seta; uropod exopod shorter than endopod articles 1–2 combined, endopod four-articled. Pseudozeuxo fischeri is characterized by pereopods 1–3 coxa with long seta about half as long as basis; pereopods 2–3 carpus with ventrodistal seta; propodus with two ventral setae; pereopods 4–6 propodus with two ventral spines and one seta; uropod endopod two- and exopod one-articled. This is the first record of the family Pseudozeuxidae and the metapseudid subfamily Metapseudinae from the Southwestern Atlantic (Brazil). Remarks on their associations with macroalgae and identification keys to world species of Apseudomorpha and Pseudozeuxo are provided.
Apseudomorpha, Brazil, new record, Rio de Janeiro, rocky shores, Tanaidomorpha
Current knowledge about the crustacean order Tanaidacea Dana, 1849 is still significantly underdeveloped, with the number of World species recognized likely to be an order of magnitude too low, with tanaidaceans potentially matching the orders Amphipoda and Isopoda in diversity mainly in environments as deep waters (
Among coastal environments, rocky shores stand out for the high biodiversity of macroalgae, invertebrates, fish and seabirds, in addition to great ecological and economic importance (
The present study is part of the Project Rocky Shores: ecology, impacts, and conservation in the areas of Região dos Lagos and the north Fluminense, which aims to survey biodiversity, study the abundance and structure of invertebrate macrofauna communities and their correlations with environmental parameters in rocky shores of the northern part of Rio de Janeiro (Macaé, Rio das Ostras and Armação dos Búzios).
The Project Rocky Shores: ecology, impacts, and conservation in the areas of Região dos Lagos and the north Fluminense has been executed by the Institute of Biodiversity and Sustainability
Sampling was carried out at four stations: Areias Negras Beach (AN) (22°31'48.98"S, 41°55'30.00"W) at Rio das Ostras; Cavaleiros Beach (CA) (22°24'17.67"S, 41°47'42.53"W) at Macaé; Calhetas Island, Santana Archipelago (AS) (22°23'54.46"S, 41°41'42.22"W), located in front of Macaé; and two points along the Rasa Beach (B1 and B2) (22°44'2.00"S, 41°57'27.36"W) (22°44'0.42"S, 41°57'26.42"W), at Armação dos Búzios.
At each site, collections were made from three zonation strata of the eulittoral zone: upper (A), intermediate (B) and lower (C). The strata were defined according to the dominant group of organisms covering each stratum, being: Upper = green macroalgae (Chlorophyta – Ulva, Enteromorpha, Chaetomorpha), bivalves (Brachidontes spp.) and small barnacles (Chthamalus spp.); Intermediate = coralline macroalgae (Rhodophyta, e.g., Corallina, Cryptonemia, Gelidium, Jania) and large barnacles (Tetraclita stalactifera); and Lower = brown macroalgae (Ochrophyta, e.g., Sargassum, Dictyota).
Five replicates were taken in each stratum, along 30 meters of extension on each shore. Sampling was made during low tides (≤ 0.2 m). The area of each sampled unit was delimited by a quadrat of 20x20 cm (0.04 m²) and samples were scraped with spatulas.
Specimens were sorted at the Laboratório Integrado de Biologia de Vertebrados, located in
Drawings were made using a microscope Zeiss with a camera lucida and digitalized with WACOM Tablet using the program Adobe Illustrator CC 2017. The appendages were dissected using chemically sharpened tungsten-wire needles. Body length was measured from the tip of the rostrum to the tip of the pleotelson, and pereonite width at the broadest part. The length/width ratio was calculated from the measurements made in the middle length and width of an article. The measurements of cheliped articles and the morphological terminology follow that used by
All material cited herein (MNRJ 29854–29871) was stored at Museu Nacional/UFRJ and was saved during the fire in this institution in September 2018, being available for future analysis.
L:W as long as wide
TBL total body length
Stn station
Four species of Tanaidacea belonging to four different families were identified in the material collected by the Project Rocky Shores in Rio de Janeiro (Brazil). One species was from the suborder Apseudomorpha (family Metapseudidae: Apseudomorpha brasiliensis sp. nov.) and three were from the suborder Tanaidomorpha – two belonging to the superfamily Paratanaoidea (families Leptocheliidae Lang, 1973: Chondrochelia dubia (Krøyer, 1842) and Pseudozeuxidae Sieg, 1982: Pseudozeuxo fischeri sp. nov.), and one of the superfamily Tanaidoidea (family Tanaididae Nobili, 1906: Zeuxo coralensis Sieg, 1980). Herein we describe the two new species as seen below.
Order Tanaidacea Dana, 1849
Suborder Apseudomorpha Sieg, 1980
Superfamily Apseudoidea Leach, 1814
Family Metapseudidae Lang, 1970
Subfamily Metapseudinae Lang, 1970
Apseudomorpha Miller, 1940: 315.
Apseudomorpha
–
See
Apseudomorpha oahuensis Miller, 1940.
Apseudomorpha albida (Shiino, 1951); A. avicularia (Barnard, 1914); A. brasiliensis sp. nov.; A. drummi Morales-Núñez, Heard & Bird, 2019; A. fontainei Guţu, 1987; A. glebosa (Menzies, 1953); A. hirsuta (Stebbing, 1910); A. magdalenensis (Menzies, 1953); A. martinicana Guţu, 2009; A. negoescuae Guţu, 2007; A. oahuensis Miller, 1940; A. ortizi Guţu, 2006; A. timaruvia (Chilton, 1882); A. veleronis (Menzies, 1953); A. vestafricana Guţu, 2006.
Family Metapseudidae is divided into four subfamilies: Chondropodinae Guţu, 2008; Metapseudinae; Msangiinae Guţu, 2006 and Synapseudinae Guţu, 1972 (
The species Parapseudes hirsutus Stebbing, 1910 was reclassified by
The genus Apseudomorpha has a worldwide distribution, with 15 species located in tropical and temperate regions of the Pacific, Atlantic and Indian Oceans (
Holotype: Brazil • 1 ♀ ovigerous, TL 2.4 mm (
Allotype: Brazil • 1 ♂, TL 1.8 mm (
Paratypes: Brazil • 1 ♀ ovigerous and 1 ♂, dissected, TL 2.0 mm (
Female. Rostrum with rounded tubercles at base and bifurcate tip. Pleonites 2 and 5 with pleura having long distal seta. Antennule article-1 inner margin with one blunt apophysis, outer flagellum with three segments. Mandible palp article-2 and article-3 with six and nine finely penicillate setae on inner margin respectively. Maxilliped palp article-1 outer margin with seta. Pereopod-1 basis with dorso-proximal margin lacking blunt, spiniform process; carpus and propodus with two and four ventral spines, respectively. Pereopods 1–2 basis with several setae along ventral margin. Pleopods biramous, exopod and endopod each with one long penicillate seta. Uropod exopod shorter than endopod segments 1–2 combined, endopod with four segments.
Male. Cheliped propodus just wider than long, ventral margin with only three simple setae (without proximal apophysis).
Based on ovigerous ♀ holotype (
Body (Figs
Cephalothorax (Fig.
Pereon (Fig.
Pleon (Figs
Pleotelson (Figs
Antennule (Fig.
Antenna (Fig.
Mouthparts: Labrum not recovered. Mandibles (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
Cheliped (Fig.
Pereopod-1 (Fig.
Pereopod-2 (Fig.
Pereopod-3 (Fig.
Pereopod-4 (Fig.
Pereopod-5 (Fig.
Pereopod-6 (Fig.
Pleopods (Fig.
Uropod (Fig.
Adult male. Length 1.8 mm. Similar to females except in the size of the chelipeds (allotype and paratype:
Cheliped (Fig.
The name is dedicated to Brazil, the country where the species were collected.
Eulittoral zone of rocky shores at Santana Archipelago, Macaé, Rio de Janeiro, Brazil.
This species was found exclusively at eulittoral zone on rocky shores (macroalgae bank) of Santana Archipelago, Macaé, Rio de Janeiro, Brazil, Southwestern Atlantic (Fig.
In total, 33 specimens of Apseudomorpha brasiliensis sp. nov. were found in eight of 60 quadrats (13.3%) collected at Santana Archipelago. This species was most abundant in the intermediate stratum (82%), however it was also found in the lower stratum (18%), being absent in the upper stratum. The substrate of the quadrats where the species was found was covered predominantly by articulated calcareous algae (Rhodophyta), covering a surface between 50–95% (average of 81% of the area). The intermediate stratum presented mainly Rhodophyta macroalgae, although brown algae Ochrophyta also occurred. The predominant Rhodophyta genera were Corallina, Jania and Arthrocardia, with other taxa such as Gracilaria, Hypnea, Pterocladiella and Plocamium. The lower stratum showed a greater coverage of brown algae, mainly of the genera Sargassum, Padina and Colpomenia.
The new species from Brazil is more similar to those Apseudomorpha species characterized by having long setae only on the second and fifth pleonites epimera, namely A. drummi, A. fontainei, A. glebosa, and A. martinicana (
Apseudomorpha brasiliensis is different from A. glebosa by (1) pereopod-1 carpus and propodus ventral margin with two and four spines, respectively (three and five in A. glebosa); (2) pereopod-6 with two ventral spines (four in A. glebosa); and (3) each pleopod rami with long penicillate seta (only one rami with two setae in A. glebosa). The new species is distinguished from A. martinicana (1) antennule article-1 inner margin with one blunt apophysis (3–4 apophyses in A. martinicana); (2) antennule main flagellum with three segments (four in A. martinicana); (3) mandible palp with article-2 with six inner penicillate setae (three in A. martinicana); and (4) pereopods 2–3 propodus with four ventral spines (three in A. martinica).
The new species from Brazil closely resembles A. drummi by having antennule main flagellum with three segments, pereopod-1 carpus and propodus with two and four ventral spines respectively and pleopods with each rami having one long penicillate seta. The former, however, can be distinguished from A. drummi by: (1) antennule article-1 inner margin with one blunt apophysis (three apophyses in A. drummi); (2) mandible palp article-2 and article-3 with six and nine finely penicillate setae on inner margin respectively (five and eight in A. drummi); (3) pereopods 1–2 basis with several setae along ventral margin, except distally (maximum one in A. drummi); (4) uropod exopod shorter than endopod segments 1–2 combined (longer in A. drummi); and (5) uropod endopod with four segments (five in A. drummi). These comparisons and others are developed in a key to the species of Apseudomorpha, shown below.
Apseudomorpha brasiliensis is the first record of the subfamily Metapseudinae from the Southwestern Atlantic. Including the present data, the family Metapseudidae is now represented by four genera in three subfamilies in Brazil: Chondropodinae (Calozodion Gardiner, 1973 and Vestigiramus Guţu, 2009), Metapseudinae (Apseudomorpha) and Synapseudinae (Synapseudes Miller, 1940).
1 | Pereopod-1 basis dorso-proximal margin with blunt, spiniform process | A. negoescuae [Indian Ocean: Réunion Island] |
– | Pereopod-1 basis dorso-proximal margin lacking blunt, spiniform process | 2 |
2 | Pleonites 1–5 without a long seta on dorsolateral margins | A. avicularia [Southeastern Atlantic Ocean: Cape Town] |
– | At least the last pleonite with a long seta on dorsolateral margins | 3 |
3 | Pleopods absent | A. vestafricana [Eastern Atlantic Ocean: Mauritania] |
– | Pleopods present (uni- or biramous) | 4 |
4 | Only the last pleonite with a long seta on distolateral margins | 5 |
– | More than the last pleonite with a long seta on distolateral margins | 6 |
5 | Antennule first peduncular article with some dentiform processes on the proximal half of inner margin | A. ortizi [Caribbean Sea: Cuba] |
– | Antennule first peduncular article without dentiform processes on the proximal half of inner margin | A. albida [Northwest Pacific Ocean: Japan] |
6 | Last four pleonites with a long seta on distolateral margins | 7 |
– | At most the last three pleonites with a long seta on distolateral margins | 8 |
7 | Pleopods biramous, with biarticulate exopodite | A. magdalenensis [Northwest Atlantic Ocean: Mexico] |
– | Pleopods biramous, with uniarticulate exopodite (in male), or uniramous (in female) | A. veleronis [Southeast Pacific Ocean and Northeast Atlantic Ocean: Ecuador, Colombia, and Panama] |
8 | Last three pleonites with a long seta on lateral margins | A. oahuensis [Southern North Pacific Ocean: O‘ahu Island] |
– | Only two pleonites (second and fifth) with a long seta on distolateral margins | 9 |
9 | Pleopods uniramous | A. fontainei [Red Sea: Aqaba Gulf] |
– | Pleopods biramous | 10 |
10 | Pereopod-1 carpus and propodus with three and five ventral spines (= spiniform setae), respectively; one rami of pleopods with two long setae | A. glebosa [Northeast and Southeast Pacific Ocean: Mexico to Ecuador] |
– | Pereopod-1 carpus and propodus with two and four ventral spines (= spiniform setae), respectively; each rami of pleopods with one long setae | 11 |
11 | Antennule main flagellum with four short, thick segments. Mandible palp article-2 with three penicillate (= setulate) setae on inner margin. Maxilliped palp article-1 outer margin with very small seta. Pereopod-2 ventral margin of propodus with three setae | A. martinicana [Caribbean Sea: Martinique Island] |
– | Antennule main flagellum with three segments. Mandible palp article-2 with more than three penicillate (= setulate) setae on inner margin. Maxilliped palp article-1 outer margin with seta. Pereopods 2–3 propodus with four ventral spines (= spiniform setae) | 12 |
12 | Mandible palp article-2 and article-3 with five and eight finely penicillate (= setulate) setae on inner margin, respectively. Uropod exopod longer than endopod segments 1–2 combined. Uropod endopod with five segments | A. drummi [South Pacific Ocean: Mo‘orea Island (French Polynesia)] |
– | Mandible palp article-2 and article-3 with six and nine finely penicillate (= setulate) setae on inner margin, respectively. Uropod exopod shorter than endopod segments 1–2 combined. Uropod endopod with four segments | A. brasiliensis sp. nov. [Southwestern Atlantic Ocean: Brazil] |
Suborder Tanaidomorpha Sieg, 1980
Superfamily Paratanaoidea Lang, 1949
Family Pseudozeuxidae Sieg, 1982
Pseudozeuxo Sieg, 1982: 66.
Pseudozeuxo
—
Pleon consisting of five pleonites (all reduced) and pleotelson, provided with only one pair of greatly reduced pleopods. Antennule with subterminal aesthetasc. Antenna article-2 with two setae. Maxilliped endites with gustatory cusps on distal margin. Cheliped carpus with three ventral setae. Pereopods 2–3 propodus with two ventral setae. Uropods exopod one or two-articled; endopod two-articled.
Pseudozeuxo belizensis Sieg, 1982.
Pseudozeuxo belizensis Sieg, 1982; P. fischeri sp. nov.
The family Pseudozeuxidae is characterized by reduced pleon with pleonites narrower than pereonite-6 in dorsal view and pleonites 1–5 combined shorter than pereonite-6 (
Pseudozeuxo differs from Charbeitanais by (1) antennule with subterminal aesthetasc; (2) antenna article-2 with two setae; (3) maxilliped endites with gustatory cusps only on distal margin; (4) cheliped carpus with three ventral setae; (5) pereopods 2–3 propodus with two ventral setae and (6) uropod endopod two-articled (
Holotype: Brazil • 1 ♀ non-ovigerous, TL 1.3 mm (
Allotype: Brazil • 1 ♂, TL 1.0 mm (
Paratypes: Brazil • 1 ♀ ovigerous (only oostegites remained), dissected TL 1.2 mm (
Female. Pereopods 1–3 coxa with long seta about half as long as basis. Pereopods 2–3 carpus with one ventrodistal seta. Pereopods 4–6 propodus with two spines and one seta ventrally. Uropod endopod article-2 0.8 times as long as article-1; exopod one-articled.
Based on non-ovigerous ♀ holotype (
Body (Figs
Cephalothorax (Figs
Pereonites 1–6 with length ratio of 0.6:0.8:0.9:1.0:1.0:0.6; pereonites 1, 2, 3 and 6 wider than long, pereonites 3–5 as long as wide; all pereonites with pair of long dorsodistal and short lateral simple setae; pereonite-1 with another pair of minute dorsodistal setae and pereonites 4–6 with pair of long mid-lateral setae.
Pleon (Figs
Pleotelson (Figs
Antennule (Figs
Antenna (Fig.
Mouthparts: Labrum (Fig.
Maxilliped (Fig.
Cheliped (Fig.
Pereopod-1 (Fig.
Pereopod-2 (Fig.
Pereopod-3 (Fig.
Pereopod-4 (Fig.
Pereopod-5 (Fig.
Pereopod-6 (Fig.
Pleopods (Fig.
Uropod (Fig.
Named in honor of Dr. Luciano Fischer (
Eulittoral zone of rocky shores at Areias Negras Beach, Rio das Ostras, Rio de Janeiro, Brazil.
This species was found within a macroalgae bank of the eulittoral zone of rocky shores at Areias Negras Beach, Rio das Ostras, Rio de Janeiro, Brazil, Southwestern Atlantic (Fig.
Geographic distribution of Apseudomorpha brasiliensis sp. nov. (circle) and Pseudozeuxo fischeri sp. nov. (square) on the study area. A–C Areias Negras Beach, Rio das Ostras. A General view of Areias Negras Beach; B view of the rocky shore eulittoral site; C view of the quadrat sample and seaweed cover; D–F Santana Archipelago, Macaé; D General view of Santana Archipelago; E view of the rocky shore eulittoral site; F view of the quadrat sample and seaweed cover. Program: QGIS 2.16.3 SRC: EPS4326 WGS84.
A total of seven specimens were found in only two of 56 quadrats (3.6%) collected at the Areias Negras Beach. The species occurred solely in the lower stratum. In the quadrats where the species was found, the substrate was mainly covered by the brown algae Sargassum (70% of the surface) and articulated calcareous algae Rhodophyta (20–25% of the surface).
The genus Pseudozeuxo is currently monotypic and includes P. belizensis (WoRMS, 2020b). Pseudozeuxo fischeri sp. nov. presents all the diagnostic characters of the genus. It is distinguished from P. belizensis by (1) pereopods 1–3 coxa with long seta about as half as long as basis; (2) pereopods 2–3 carpus with only seta ventrodistally (P. belizensis has seta and spine); (3) pereopods 4–6 propodus with two spines and one seta ventrally; (4) uropod endopod article-2 0.8 times as long as article-1 (P. belizensis with article-2 small, about 0.4 times as long as article-1) and exopod one-articled (P. belizensis is two-articled).
Charbeitanais and Haimormus are recorded from the Pacific Ocean (Hong Kong and Japan, respectively;
1 | Pereopods 1–3 coxa with short seta (less than half of basis). Pereopods 2–3 carpus with seta and spine ventrodistally. Pereopods 4–6 propodus with two setae ventrally. Uropod endopod article-2 small, about 0.4 times as long as article-1, and exopod two-articled | P. belizensis [Caribbean Sea: Belize] |
– | Pereopods 1–3 coxa with long seta (about half as long as basis). Pereopods 2–3 carpus with only seta ventrodistally. Pereopods 4–6 propodus with two spines and seta ventrally. Uropod endopod article-2 0.8 times as long as article-1 and exopod one-articled | P. fischeri sp. nov. [Southwestern Atlantic Ocean: Brazil] |
The apseudomorphs are predominantly a shallow-water group and the family Metapseudidae occurs mainly in the continental shelf <200 m (
Currently, there are regions whose shallow-water tanaidacean faunas are still under-researched. A study of the Tanaidacea from Australia, predominantly in shallow waters, has discovered an extremely high diversity, increasing the number of described species in 14 years from 16 in 1996 to 117 by 2010 (
Apart from that, six from the 54 species registered in Brazilian coast were recorded in sediments with algae (Apseudes aisoe Araújo-Silva, Coelho and Larsen, 2013, A. noronhensis Araújo-Silva, Coelho and Larsen, 2013, Chondrochelia dubia, Intermedichelia jesseri Araújo-Silva & Larsen, 2012, Makraleptochelia potiguara Araújo-Silva & Larsen, 2012, Zeuxo coralensis), representing three families: Apseudidae Leach, 1814, Leptocheliidae and Tanaididae (Brum 1973;
The diversity of the tanaidaceans in certain ocean regions remains to be discovered, especially in Brazil where this group is still understudied (in rocky intertidal sites, shallow water sediment or deep sea environments). Both new species represent also new records to Southwestern Atlantic for the family Pseudozeuxidae (Pseudozeuxo fischeri sp. nov.) and for the subfamily Metapseudidae (Apseudomorpha brasiliensis sp. nov.). Thus, it is likely that further studies on the Brazilian coast would lead to the discovery of new species besides other new records.
Great advances have been made in the past twenty years to understand the Brazilian Tanaidacean fauna; however progress will continue to be restricted by the lack of tanaidacean taxonomists and researchers, but mainly by the lack of funding for this enterprise. Nevertheless, it is evident from our current knowledge that the Tanaidacea form a very diverse order of the Peracarida and are of considerable ecological significance in certain regions and habitats (
This research was carried out within the scope of the “Project Rocky Shores/FUNBIO”, and was supported by an environmental offset measure established through a Consent Decree/Conduct Adjustment Agreement between Chevron Brazil and the Brazilian Ministry for the Environment, with the Brazilian Biodiversity Fund – FUNBIO as implementer. The authors are grateful to Dr. Luciano Fischer (