Checklist and new records of Lophogastrida (Malacostraca, Peracarida) in Chilean waters

Abstract

Specimens of lophogastrids were collected off Chile, including four first records: Ceratolepis hamata G.O. Sars, 1883 in the vicinity of Easter Island, Eucopia unguiculata (Willemoës-Suhm, 1875) the Juan Fernández Archipelago, Gnathophausia zoea Willemoës-Suhm, 1873 off Concepción, and Neognathophausia ingens (Dohrn, 1870) off Valparaíso. Additional records were also obtained for Chile for four species: Eucopia australis Dana, 1852, Eucopia grimaldii Nouvel, 1942, Fagegnathophausia gracilis (Willemoës-Suhm, 1875), and Paralophogaster sp. Based on these records, lophogastrids are now represented off Chile, in the southeastern Pacific, by nine species.

Keywords:
Biodiversity; Chile; Lophogastrida; Peracarida; southeastern Pacific

INTRODUCTION

The order Lophogastrida is little known worldwide and is considered more primitive than its closest and better-studied relative order Mysida. Species of Lophogastrida are considered of great importance in the links for energy conversion in the oceans, and they are considered as one of the main consumers of zooplankton in deep water. The scarcity of studies is in great part related to their habitat and the difficulty of routine sampling in deep waters (Meland and Willassen, 2007; Meland et al., 2015).

Species of Lophogastrida inhabit both meso- and bathypelagic environments in the oceans worldwide (Castellani et al., 2017). They belong to three families: Lophogastridae G.O. Sars, 1870, Eucopiidae G.O. Sars, 1885, and Gnathophausiidae Udrescu, 1984 (Meland and Willassen, 2007). According to Clarke (1962), the oldest record of a lophogastrid (group that gave rise to the GnathophausiaWillemoës-Suhm, 1873) was that of M. Sars who reported, in 1857, Lophogaster typicus in his report of the “Challenger” campaigns carried out between 1873 and 1876. Years later, G.O. Sars (1885) recognized nine species in the genus Gnathophausia from different parts of the world (Ibarra and Hendrickx, 2008).

Some of the most complete works dealing with the eastern Pacific lophogastrids are those by Tattersall (1951) and Banner (1954), who studied the material from the “Americas” included in the collections of the Smithsonian Institution, Washington D.C. Additional records of species, or new material of lophogastrids, were also reported by Austin (1985), Brusca and Hendrickx (2005), Ibarra and Hendrickx (2008), Hendrickx (2019), Hendrickx and López (2019), and Huber et al. (2019). A complete checklist of lophogastrids occurring in the Mexican Pacific was proposed by Hernández-Payán and Hendrickx (2020) who reported eight species from this area. Three families of Lophogastrida are represented in Chilean waters. Lophogastrida differs notably from the Mysida, with the major differences being well-developed gills on most thoracopods for both sexes, the presence of pleopods in both sexes, a marsupium consisting of seven pairs of oostegites in females, and the absence of a statocyst in the external branch of the uropod (Nouvel et al., 1999), although some species of deep-sea Mysida, like the family Petalophthalmidae, also lack statocysts in the base of uropods and have oostegites. Pleural plates are present in the Lophogastridae and Gnathophausiidae and absent in Eucopiidae (except in EucopiaDana, 1852). The Gnathophausiidae feature a long and thin rostrum, armed with small teeth, while in the Lophogastridae the rostrum is unarmed, short, and wide (Casanova et al., 1998; Murano, 1999).

Neognathophausia gigas (Willemoës-Suhm, 1875) and Fagegnathophausia gracilis (Willemoës-Suhm, 1875) (both previously belonged in the genus Gnathophausia) have been cited for the Chile-Peru and Iquique trench, respectively (Soto and Retamal, 1991). Brandt et al. (1998) cited Eucopia australisDana, 1852 from the southern tip of South America.

Studies carried out by the former Soviet Union in the region indicate the presence of five species of lophogastrids in the vicinity of the seamounts of the Salas and Gómez and the Nazca mountain ranges (Vereshchaka, 1990).

During biodiversity studies of the Chilean mesopelagic zone, specimens of nine species of lophogastrids were collected, none of which had been previously recorded from the southeastern Pacific (SEP). This material is reported herein and includes new records for the Chilean fauna.

MATERIAL AND METHODS

The specimens reported here were collected by different expeditions off the coast of Chile. In the northern zone, material was obtained from the United Nations development project (FAO-UNDP CHI/87/007) between longitude 18º38'S and 21º47'S and from the coast to latitude 72º04'W. Other material was collected by the CIMAR programs of the National Oceanographic Committee expeditions to the Chilean Oceanic Islands (CIMAR 5, CIMAR 6, CIMAR 21, CIMAR 22), and to the Chilean fjords (CIMAR 14, CIMAR 25). Other expeditions included the MOPEX Expedition off Valparaiso, the ONR expedition off Concepción, and the ATACAMEX expedition in the Atacama trench off Antofagasta. Specimens were collected with an IKMT (Isaacs-Kidd Midwater Trawl) net. The material examined is deposited in the Museum of the Sea of the Arturo Prat University. A restricted synonymy is provided for each species, including the prime synonymy and references relevant to this study.

RESULTS

Nine species were registered during this study, four of which are recorded for the first time in waters of the SEP off Chile:

Order Lophogastrida Boas, 1883

Family Eucopiidae G.O. Sars, 1885

Genus Eucopia Dana, 1852

Eucopia australis Dana, 1852

Eucopia australisDana, 1852: 609-611, pl. 40 (10a-m). - Hansen, 1913: 8. pl. I, fig. 1a, b. - Illig, 1930: 404‒405. - Fage, 1942: 41‒47, figs. 28a, 29‒35. - Tattersall, 1951: 33 [distribution, remarks]. - Tattersall, 1955: 48‒49, fig. 4C, D. - Birstein and Tchindonova, 1962: 61. - Kathman et al., 1986: 140‒141. - Wittmann, 1990: 130, fig. 257.

Figure 1.
Localities where species of the order Lophogastridae are present in Chilean waters.

Material examined. 3 females, station E-14 PNUD/Chi-87 (20°20'S 70°58'W) (Fig. 1).

Previous records from SEP. This is the first record from the SEP.

Geographic distribution. Cosmopolitan, meso- to bathypelagic, under 600 m depth (Mauchline and Murano, 1977).

Depth range. 600‒6,050 m (Müller, 1993).

Eucopia grimaldii Nouvel, 1942

Eucopia grimaldiiNouvel, 1942: 5, 6, figs. 5, 6.

Eucopia grimaldii - Birstein and Tchindonova, 1962: 61, 62. - Kathman et al., 1986: 144, 145.

Eucopia grimaldi - Vereshchaka, 1990: 119.

Material examined. 2 females, station 4 CIMAR 21 (26º59'54''S 71º12'57''W).

Previous records from SEP. In the vicinity of the “Soldier” seamount (21º41'S 81º46'W) of the Salas & Gomez submarine ridge (Vereshchaka, 1990).

Geographic distribution. Pan oceanic from Alaska to the Antarctic (San Vicente, 2016).

Depth range. 300-5,430 m (most commonly in about 2,000 m) (Müller, 1993; Wittman, 2020a).

Eucopia unguiculata ( Willemoës-Suhm, 1875 )

Chalaraspis unguiculataWillemoës-Suhm, 1875: 40, pl. 8.

Eucopia unguiculata. - Hansen, 1905: 3. - Hansen, 1910: 20, 21, pl. I, fig. 3a. - Hansen, 1912: 187-188; - Hansen, 1927: 20. - Holt and W.M. Tattersall, 1906: 20 (and references therein). - Tattersall, 1951: 34 [distribution]. - Birstein and Tchindonova, 1962: 62. - Austin, 1985: 560. - Kathman et al., 1986 (and references therein): 145, 147, text figs. a, b, d, f, h. - Price et al., 2009: 927.

Material examined. 1 male, station 26 CIMAR 22 (33º45'00''S 81º00'00''W); 1 male, station 34 CIMAR 22 (33º36'00''S 79º15'00''W); 2 females and one male, station 38 CIMAR 22 (33º00'00''S 73º00'00''W) (Fig. 1).

Previous records from SEP. First record from the Southeast Pacific.

Geographic distribution. Pan oceanic in tropical to Arctic waters, latitudes all along from Arctic to Antarctic oceans (Wittmann, 2020 b ).

Depth range. 100-6,500 m (Müller, 1993).

Family Gnathophausiidae Udrescu, 1984

Genus Fagegnathophausia Petryashov, 2015

Fagegnathophausia gracilis ( Willemoës-Suhm, 1875 )

Gnathophausia gracilisWillemoës-Suhm, 1875: 33, pl. 9, fig. 1.

Gnathophausia gracilis - G.O. Sars, 1885: 48, pl. VII, figs. 6-10. - Ortmann, 1907: 39. - Hansen, 1912: 185, 186. - Fage, 1941: 27, fig. 27. - Tattersall, 1951: 28. - Pequegnat, 1965: 409, fig. 6. - Vereshchaka, 1990: 119.

Gnathophausia brevispinis - Faxon, 1895: 216, pl. J.

Gnathophausia dentataFaxon, 1893: 217.

Material examined. 1 male associated with the "centollón del norte" (Lithodidae) fishing offshore of Iquique in 2001; 1 male collected during the ATACAMEX expedition in 2018 to the Atacama trench.

Previous records from SEP. In the vicinity of the “Soldier” seamount (21º41'S 81º46'W).

Geographic distribution. Widely distributed from 34º N to 22ºS, bathypelagic (Mauchline and Murano, 1977; Soto and Retamal, 1991; Murano, 1999).

Depth range. 500-5,225 m (Müller, 1993; Wittmann, 2020 a ).

Gnathophausia zoea Willemoës-Suhm, 1873

Gnathophausia zoeaWillemoës-Suhm, 1875: 32, pl. 9, figs. 2-15. - G.O. Sars, 1885: 44. - Faxon, 1896: 164. - Ortmann, 1907: 42, pl. 2, fig. 2a, b. - Hansen, 1910: 17; - Hansen, 1912: 186. - Fage, 1941: 34. - Tattersall, 1951: 29. - Birstein and Tchindonova, 1958: 261. - Pequegnat, 1965: 410. - Vereshchaka, 1990: 119. - Brusca and Hendrickx, 2005: 141.

Gnathophausia Willemoësii G.O. Sars, 1885: 38, pl. 5, figs. 1-6. - Faxon, 1895: 215, pl. K, fig. 1. - Ortmann, 1906: 969.

Gnathophausia zoea sarsi - Ortmann, 1907: 42.

Material examined. 1 specimen collected by the expedition ONR agt11 in front of Concepción (32º21'S 73º43'W) (Fig. 1).

Previous records from SEP. First record from the Southeast Pacific.

Geographic distribution. Cosmopolitan in temperate and tropical zones of the Pacific, Indian and Atlantic oceans, between 65°N‒37°S (Mauchline and Murano 1977; Lagardere, 1983; Brusca and Hendrickx, 2005; Hendrickx, 2019).

Depth range. 400-6,050 m (Müller, 1993).

Neognathophausia ingens ( Dohrn, 1870 )

Lophogaster ingensDohrn, 1870: 610, pl. 31, figs. 12-14.

Gnathophausia ingens - G.O. Sars, 1885: 30, pl. II. - Ortmann, 1907: 28, pl. II, fig. 1a. - Hansen, 1912: 184, 185. - Fage, 1941: 15. -Tattersall, 1951: 25. - Clarke, 1961: 313, fig. 1. - Birstein and Tchindonova, 1962: 59. - Pequegnat, 1965: 403, figs. 3, 4. - Kathman et al., 1986: 160. - Escobar-Briones, 2002: 295.

Gnathophausia calcarata G.O. Sars, 1885: 35, pl. IV. - Ortmann, 1907: 30, pl. 1, fig. 2a-f.

Neognathophausia ingens - Petryashov, 1992: 47, 48.

Material examined. 1 specimen damaged, station 38 CIMAR 22 (33º00'00''S 73º00'00''W) (Fig. 1).

Previous records from SEP. First record from the Southeast Pacific.

Geographic distribution. Cosmopolitan, between 40°N and 45°S (Mauchline and Murano, 1977).

Depth range. 225-3,914 m (Müller, 1993).

Family Lophogastridae G.O. Sars, 1870

Genus Ceratolepis G.O. Sars, 1883

Ceratolepis hamata G.O. Sars, 1883

Ceratolepis hamata G.O. Sars, 1883: 4.

Ceratolepis hamata - Fage, 1941: 9-14, figs. 11-19. - Vereshchaka, 1990: 118.

Material examined. 4 females, station e-43 CIMAR-5 (27º00'30''S 109º12'17''W).

Previous records from SEP. In the vicinity of the seamounts “Rock” (25º58'S 100º41'W), “Ichthyologist” (25º07'S 99º35'W) and “Dome” (25º04'S 97º26'W) (Vereshchaka, 1990).

Geographic distribution. Samoa, Indo-Pacific between 10º and 25ºS, epi- and mesopelagic between 25 and 300 m (Mauchline and Murano, 1977).

Depth range. ?25-4,780 m (Müller, 1993).

Genus Paralophogaster Hansen, 1910

Paralophogaster glaber Hansen, 1910

Paralophogaster glaberHansen, 1910: 16, 17, pl. I, fig. 2a-n. - Tattersall, 1951: 22-24 [distribution, remarks].

Paralophogaster glaber pacificusVereshchaka, 1990: 119-122, fig. 1.

Material examined. 1 female and 2 males, station E-74 CIMAR 5 (26º28'06''S 105º10'45''W). In the vicinity of Salas and Gómez Islands. (Fig. 1).

Previous records from SEP. First record from the Southeast Pacific.

Geographical distribution. West Pacific Atlantic between 40ºN and 35ºS (Tattersall, 1951; Mauchline and Murano, 1977).

Depth range. Surface to 310 m (Müller, 1993).

DISCUSSION

Lophogastrids correspond to non-decapod pelagic organisms similar to shrimp, mostly with distributions limited to deep waters. Their records seem to be accidental in plankton samples, for which special sampling equipment for deep waters is required. In Chile they have been reported occasionally (Soto and Retamal, 1991).

The genus Eucopia contains nine valid species (Mees and Meland, 2012), including one fossil species (Eucopia praecursorSecretan and Riou, 1986†). Most species of this genus have a wide distribution and several are cosmopolitan (Kathman et al., 1986; Burghart et al., 2007).

Eucopia australis has been cited for the SEP; however, the group of lophogastrids in the SEP present complexities in the taxonomic status of some of them. Three species of Eucopia have been recorded in the SEP, but the few records available do not allow establishment of distributional patterns in this area of the Pacific. Of these three species, E. grimaldii has been recorded in the vicinity of seamounts by Vereschaka (1990). The differences between the three species are relatively clear. The telson of E. unguiculata has a very truncate apex armed with two long spines, this character is unique to this species. In the case of E. australis and E. grimaldii, they can be differentiated by a series of characters such as the shape of the anterior margin of the carapace; i.e., broad in E. grimaldii and narrow in E. australis, the relationship between the length of the sixth pleonite and the length of the telson, and the shape of the end of the telson and its armature.

Species of Gnathophausidae are quite characteristic for their intense red color and their body ornamented with thorns and keels. The taxonomic status of the species of this family has been reviewed rather extensively and there is clarity in this regard (Meland and Aas, 2013; Petryashov, 2015).

The family Lophogastridae is represented in the SEP by three species or subspecies, C. hamata, P. glaber glaber, and P. glaber pacificus. Ceratolepis hamata has been collected in the SEP only on two occasions, both in the vicinity of the seamounts of the Salas and Gómez Mountain ranges (Vereshchaka, 1990; this study), and this record of C. hamata would be the westernmost limit of this species. Both subspecies of P. glaber are present in the SEP. According to Vereshchaka (1990), the specimens collected in the vicinity of the seamounts of the Salas and Gómez Mountain range differ from the typical P. glaber, something that had already been noted by Bacescu (1981) in specimens from different locations analyzed in Bacescu (1981). The specimens recorded here match the original description of Hansen (1910) and differ from that indicated by Vereshchaka (1990). The number of spines on the telson corresponds well with the original description of Hansen and that indicated by other authors (Hansen, 1910; Tattersall, 1951; Bacescu, 1981).

In the Northeast Pacific, eight species of the order Lophogastrida have been cited (Ibarra and Hendrickx, 2008; Hendrickx, 2019; Hernández-Payán and Hendrickx, 2020) of which four are present in the SEP: E. unguiculata, F. gracilis, G. zoea, and N. ingens.

Hernández-Payán and Hendrickx (2020), taking into account the information reported by Austin (1985), Brusca and Hendrickx (2005), Ibarra and Hendrickx (2008), Mees and Meland (2012), San Vicente (2016), Hendrickx (2019), and Hernández-Payán (2020), revaluated the fauna of Lophogastrida for the eastern Pacific (from Alaska to Chile) and concluded that there are 13 species of Lophogastrida known from this region. Based on this study, two species are added to this list (E. unguiculata, Eucopia sculpticaudaFaxon, 1893) thus reaching a total of 15 species currently know from the eastern Pacific (Tab. 1).

Table 1.
Updated list of Lophogastrida species recorded in eastern Pacific waters (Alaska to Chile). Species reported in the Chilean Pacific in bold. FR = first record in Chilean waters.

Order Lophogastrida
Family Eucopiidae G.O. Sars, 1885
Genus EucopiaDana, 1852
Eucopia australisDana, 1852 FR
Eucopia grimaldi Nouvel, 1942
Eucopia major Hansen, 1910
Eucopia sculpticauda Faxon, 1893
Eucopia unguiculata (Willemoës-Suhm, 1875) FR
Family Gnathophausiidae Udrescu, 1984
Genus FagegnathophausiaPetryashov, 2015
Fagegnathophausia gracilis (Willemoës-Suhm, 1875)
Genus GnathophausiaWillemoës-Suhm, 1873
Gnathophausia childressi Casanova, 1996
Gnathophausia scapularis Ortmann, 1906
Gnathophausia zoeaWillemoës-Suhm, 1873 FR
Genus NeognathophausiaPetryashov, 1992
Neognathophausia gigas (Willemoës-Suhm, 1873)
Neognathophausia ingens (Dohrn, 1870) FR
Family Lophogastridae G.O. Sars, 1870
Genus Ceratolepis G.O. Sars, 1883
Ceratolepis hamata G.O.Sars, 1883
Genus Chalaraspidum Willemoës-Suhm, 1895
Chalaraspidum alatum (Willemoës-Suhm, 1876)
Genus Lophogaster M. Sars, 1857
Lophogaster japonicus W.M. Tattersall, 1951
Genus ParalophogasterHansen, 1910
Paralophogaster glaber Hansen, 1910

Recent studies of deepwater have increased the number of species of the order Lophogastrida in Chilean waters (Fig. 1). However, a greater number of samples are required in the area as well as the use of specialized equipment to obtain stratified samples to determine the vertical distribution of the different species. It is also recommended to obtain new material not preserved in formaldehyde for genetic analysis to clarify the phylogenetic relationships among species.

ACKNOWLEDGMENTS

Thanks go to the CIMAR Ocean Islands Program of CONA, and to Dr. Rubén Escribano of the Millennium Institute of Oceanography. Dr. V. A. Gallardo is thanked for providing material from the ONR Expedition in front of Concepción and to the Vice-Rectory of Research, Innovation and Postgraduate Studies of the Arturo Prat University for the financing of the VRIIP0093-17 Project.

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• Author Contributions

  Conceptualization and Design: JCH-P. Performed research: JCH-P, GG, NOC. Acquisition of data: JCH-P, GG, NOC. Analysis and intepretation of data: JCH-P, GG, NOC. Writing - original draft: JCH-P, GG, NOC. Writing - critical review & editing: JCH-P, GG, NOC.

• Consent for publication

  All the authors give their consent, approve, and give the go-ahead for the publication of this manuscript in their journal.

• Competing interests

  There is no conflict of interest between the authors of this manuscript.

• Data availability

  There is no problem for the availability of the data, the material used for this manuscript is deposited in museum collections.

• Funding and grant disclosures

  The material reported in this study was obtained thanks to the financing of several projects, one of them was from the United Nations development project (FAO-UNDP CHI/87/007), another material was collected by the CIMAR programs from the Committee's expeditions. Oceanografico Nacional de las Islas Oceanicas de Chile (CIMAR 5, CIMAR 6, CIMAR 21, CIMAR 22) in addition to the Chilean fjords (CIMAR 14, CIMAR 25), also the material collected was obtained from the financing of the project VRIIP0093-17 by the Vice-Rector for Research, Innovation and Postgraduate Studies at the Arturo Prat University.

• Study Association

  This study was carried out in association with different institutions and with the financing of different projects.

• Study permits

  All necessary testing permits will be provided for the collection of material used in this research work.

• Zoobank:

  http://zoobank.org/urn:lsid:zoobank.org:pub:76AC8DCE-01FB-4202-B774-3330CC77CA8C

Edited by

Associate Editor:

Shane Ahyong

Data availability

There is no problem for the availability of the data, the material used for this manuscript is deposited in museum collections.

Publication Dates

• Publication in this collection
  10 Nov 2023
• Date of issue
  2023

History

• Received
  14 Nov 2022
• Accepted
  27 Apr 2023