4121 (1): 049 058 Article · 1Department of Invertebrate Zoology, Biological Faculty, Lomonosov...

ZOOTAXA

ISSN 1175-5326 (print edition)

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Zootaxa 4121 (1): 049–058

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http://doi.org/10.11646/zootaxa.4121.1.2

http://zoobank.org/urn:lsid:zoobank.org:pub:81B43683-DF38-4595-BFA8-0D4DF9E7E426

A new subspecies of Ovalona setulosa (Megard, 1967) (Cladocera: Anomopoda:

Chydoridae) from the Caribbean coast of Colombia

ARTEM Y. SINEV1,2 & JUAN M. FUENTES-REINÉS3

1Department of Invertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Leninskie Gory, Moscow 119991, Rus-

sia 2A. N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071, Russia. E-mail: [email protected] del Magdalena, Grupo de investigación en Biodiversidad y Ecología Aplicada. A.A 731 Santa Marta, Magdalena,

Colombia. E-mail: [email protected]

Abstract

A new subspecies of Ovalona setulosa (Megard, 1967) (Cladocera: Anomopoda: Chydoridae) is described based on ma-

terial from Laguna Navío Quebrado (La Guajira Department), Colombia. Ovalona setulosa arangureni ssp. nov. differs

from the nominative subspecies in: (1) major head pores position on head shield; (2) posterior margin of labral keel with-

out clusters of short setules; (3) more elongated postabdomen, with length/height ratio about 2.7-3; (4) male postabdomen

with straight dorsal margin in whole distal portion. In outer morphology and morphology of postabdomen, O. setulosa

arangureni ssp. nov. can be confused with O. glabra (Sars, 1901), but, like nominative subspecies, differs from it by in-

terrupted connection between major head pores. Our data suggest that diversity of Aloninae species with limited distribu-

tion areas in South America is underestimated yet, and areas north and south of Amazon basin can have different local

species.

Key words: Cladocera, Anomopoda, taxonomy, new subspecies, South America

Introduction

Species historically assembled into the genus Alona Baird, 1843 form the largest and most diverse assemblage within the family Chydoridae; it comprises over 80 valid species worldwide which represent about 17 % of the total species number in Cladocera (Van Damme et al. 2010). Alona s. lato is an artificial assemblage, according to Van Damme & Dumont (2008a), being Alona s. str. a small group, formed by the type species of the genus Alona

quadrangularis (O.F. Mueller, 1775) and its siblings. Alona s. lato is composed of several species-groups of generic status. Some of these groups already attained generic status, for example rectangula-group was placed into the genus Coronatella Dybowski & Grochowski, 1894 (Van Damme & Dumont 2008b), verrucosa-group—into the genus Anthalona Van Damme, Sinev & Dumont, 2011 (Van Damme et al., 2011b), pulchella-group—into genus Ovalona Van Damme & Dumont 2008 (Sinev 2015). Several more species-groups—affinis, costata, guttata

and elegans are not yet formally recognized as genera and await revision.Twenty species of Alona s. lato had been reported to Neotropic (Kotov et al. 2013), but diversity of the group

in the region is probably much higher. For example, recent studies of Alona s. lato from Central Mexican Plateau revealed two new species of the group (Sinev & Silva-Briano 2012), and revision of Coronatella from Brazil revealed three new species (Sousa et al. 2015b).

In Colombia, the knowledge of Alona s. lato still remains inadequate. Stingelin (1913) reported four species and one variety from Colombia Mountains: A. pulchella King, 1853, A. affinis (Leydig, 1860), A. costata Sars, 1862, A. guttata Sars, 1862, and A. guttata var. tuberculata. According to modern data, A. affinis and A. pulchella

(now a member of Ovalona) are not present in America (Sinev, 1998, 2001a, 2015), being substituted here by the sibling-species Alona ossiani Sinev, 1998 and Ovalona glabra (Sars, 1901), respectively, and Stingelin's (1913) report probably refers to these species. Alona costata is a Holarctic species, and it is not present in Neotropics as

Accepted by M. Alonso: 2 May 2016; published: 7 Jun. 2016 49

well. Three other species of costata-group, A. iheringula Kotov & Sinev, 2004 (= A. iheringi Sars, 1901), A. hudeci

Sinev, 1999 (= Alona rusticoides Hudec, 1990), and A. margipluma Sousa, Santos, Güntzel, Diniz, de Melo Junior & Elmoor-Loureiro, 2015 are present in South America (Sinev, 1999, 2001b, Kotov & Sinev, 2004, Sousa et al., 2015c), but it is impossible to attribute Stingelin (1913) report of A. costata to any of these taxa. A. guttata s. str. is considered a Palaearctic species, but recent studies species revealed no differences in morphology of parthenogenetic female between European and Mexican populations of the species (Sinev & Silva-Briano, 2012). According to modern data, tuberculated varieties of Alona s. lato, including A. guttata var. tuberculata, cannot be treated as a separate subspecies, once sculpture of valves is subjected to intrapopulational variability (Van Damme et al., 2010).

Investigations of Colombian Alona s. lato were continued only recently. Fuentes-Reinés et al. (2012) confirmed the presence of O. glabra and found one more species, Alona dentifera Sars, 1901 in material from Ciénaga Grande de Santa Marta, Magdalena Department (the taxonomic status of this species is unclear, Chatterjee et al. (2013) place it within gthe genus Coronatella). Fuentes-Reinés (2014) recorded O. glabra from La Guajira Department, however a detailed re-examination of this material led us to conclude that it belongs to a new taxon rather than O. glabra. The aim of this paper is to describe this new form and define its taxonomic status.

Material and methods

Plankton samples were taken from Laguna Navío Quebrado monthly between April and December 2012, mainly in the macrophyte zone, but also in the open water using a bucket of 25 L at both vegetation areas and shallow open water. All samples were filtered through a zooplankton net (45 µm) and preserved in 70% ethanol. In the laboratory, specimens were selected from samples under a binocular stereoscopic microscope, and studied under a compound optical microscope in a drop of a glycerol-formaldehyde mixture in toto. Images were taken using a Kodak Easy Share C140 digital camera attached to a compound Olympus CX22 microscope. The specimens were measured in lateral position, from the anterior end of the rostrum to the posterior margin of the valve. The specimens examined were deposited at the Museo de Colecciones Biológicas at the Universidad del Atlántico (UARC), Colombia.

Results

Order Anomopoda Sars, 1865

Family Chydoridae Dybowski & Grochowski, 1894 emend. Frey, 1967

Subfamily Aloninae Dybowski & Grochowski, 1894 emend. Frey, 1967.

Tribe Alonini Dybowski & Grochowski, 1894 emend. Kotov, 2000

Genus Ovalona Van Damme & Dumont, 2008 emend. Sinev, 2015

Alona setulosa arangureni ssp. nov.

(Figs. 1–4)

Etymology. The new subspecies is named after the Colombian researcher Dr. Nelson Aranguren, for his extensive work on the zooplankton from aquatic systems of Colombia and for his legacy and leadership of new generations of planktologists.

Type locality. Laguna Navio Quebrado, La Guajira Department, Colombia (11º25’N and 73º5’W). From this locality, 30 parthenogenetic females, 3 ephippial females, and 20 adult males were collected in June and October of 2012 by J. M. Fuentes-Reinés.

Type material. Holotype: parthenogenetic female deposited at the Museo de Colecciones Biológicas de la Universidad del Atlántico, Colombia, UARC246M.

SINEV & FUENTES-REINÉS50 · Zootaxa 4121 (1) © 2016 Magnolia Press

FIGURE 1. Alona setulosa arangureni ssp. nov. from Laguna Navio Quebrado, Colombia (type locality). A–J,

parthenogenetic female: A, lateral view; B, head shield; C, head pores; D, variants of main head pores morphology; E, labrum;

F, postabdomen; G, postanal margin of postabdomen; H, postabdominal claw; I, inner distal lobe (IDL) of limb I; J, inner

portion of limb IV. K–L, adult male: K, outline of the body; L, postabdomen.

Allotype: adult male from type locality, UARC247M. Paratypes: 5 undissected females in alcohol, UARC248M; 1 dissected female, UARC243M-257M, 3

undissected ephippial females in alcohol, UARC249M; 3 adult males, UARC250M; 1 dissected male, UARC251M-252M.

Description. Parthenogenetic female. In lateral view, body ovoid (Fig. 1A, 2A), maximum height in middle, height/length about 0.66–0.76 in adults. Dorsal margin of valves generally uniformly curved, postero-dorsal angle smoothed, postero-ventral angle broadly rounded. Ventral margin of valves with 40–45 setae of different size in different regions of the margin. Posteroventral angle of valves with about 100 short setules of similar length, not differentiated into groups. Valves oblique, without any sculpture.

Head small, not keeled, rostrum short and blunt. Compound eye and ocellus subequal in size, distance from tip of rostrum to ocellus somewhat larger than between ocellus and eye.

Head shield elongated, length/width ratio about 1.5–1.6, rostrum short, broadly rounded, posterior portion widely rounded, with wavy margin (Fig. 1B, 2B). Three major head pores of equal size, with interrupted connection between them (Fig. 1C–D), distance from posterior pore to posterior margin of head shield (PP) about

Zootaxa 4121 (1) © 2016 Magnolia Press · 51A NEW SUBSPECIES OF OVALONA SETULOSA

0.4–0.5 distance between anterior and posterior pores (IP). Lateral head pores minute, located about 1.1–1.5 IP from midline, at the level of anterior major head pore.

FIGURE 2. Alona setulosa arangureni ssp. nov. from Laguna Navio Quebrado, Colombia (type locality), parthenogenetic

female. A, adult parthenogenetic female, general view. B, head shield. C, labrum. D, postabdomen. E, antennule. F, antenna.

Labrum (Fig. 1E, 2C): distal labral plate without setulation. Labral keel moderately wide, with height about 1.3 widths. Anterior margin convex, apex rounded or blunt, posterior margin convex, without clusters of short setules.

Postabdomen (Fig. 1F, 2D) with almost parallel margins, moderately wide, length/width ratio about 2.5–2.6. Ventral margin straight or weakly convex. Dorsal margin convex in preanal part, concave in anal part, straight in postanal part. Distal margin almost straight; dorso-distal angle weakly prominent, almost right, with rounded tip.


Preanal angle well-expressed, postanal angle weakly defined. Distal portion 1.5 times longer than preanal margin; anal and postanal portions subequal in length. Postanal margin provided with 7–8 marginal denticles; anal margin with 4–5 clusters of setulae (Fig. 1G). Each postanal denticle with 1–3 short spines near the base. Length of postanal denticles evenly decreases basally; length of distalmost denticle slightly less than width of postabdominal claw base. Postanal portion with 5–6 wide, closely spaced lateral fascicles of long setulae; distalmost setule in each fascicle being the longest and thickest, 1.5 times longer than neighboring marginal denticles. Postabdominal claw(Fig. 1H) regularly curved, somewhat longer than preanal portion of postabdomen, with fine setulation along ventral margin. Basal spine length about 0.33 of claw length.

Antennule (Fig. 2E) with four clusters of setules on inner face. Antennular sensory seta slender, arising at 2/3 distance from the base. Nine aesthetascs, the longest of them about 0.8 length of antennule. All aesthetascs projecting beyond anterior margin of the head shield.

Antenna of moderate size (Fig. 2F). Antennal formula: setae 0-0-3/1-1-3; spines 1-0-1/0-0-1. In both branches, length of segments decreasing distally. Seta arising from basal segment of endopodite thin, reaching to the end of endopodite. Seta arising from middle segment of endopodite of same size with apical setae. Basal and middle segments of endopodite with clusters of hard setules. Apical setae of both branches of similar thickness. Spine on basal segment of exopodite shorter than middle segment. Apical spines longer than apical segments. Mandible

elongated; distal portion with sharp denticles (Fig. 3A)Thoracic limbs: five pairs.Limb I (Fig. 1I, 3B–D). Epipodite oval. ODL seta longer than IDL setae. IDL with three setae; seta 1 short;

setae 2–3 long, of similar length, armed with thin setulae in distal portion. Endite 3 with four setae subequal in length. Endite 2 with three setae of different length, middle one (e) longer than ODL seta. Endite 1 with two 2-segmented setae (g–h), both setulated in distal part, without a flat plumose seta shifted to the limb base. Fascicles of thin setulae on inner face of limb, plus bunches of longer and more robust setulae at ventral margin of limb. Two unequal ejector hooks.

Limb II (Fig. 3E) exopodite narrow elongated lobe with slender seta of half length of exopodite. Inner portion of limb (“endopodite”) with eight scraping spines; scrapers 1–4 long, increasing progressively in length distally; scraper 5 longer than 4; scrapers 6–8 short, subequal in length. Distal armature of gnathobase with four elements. Filter plate II with seven setae, the posteriormost member considerably shorter than others.

Limb III (Fig. 3F). Epipodite oval, without projection. Exopodite with seven setae, seta 3 being longest (Fig. 6D). Distal endite with 3 setae (Fig. 6E); distalmost and middle setae of same size, slender and sharp, with denticles in distal part; basalmost seta short, geniculated, with thin setulae. Basal endite with 4 stiff, feathered in distal part setae, increasing in size in basal direction. Gnathobase filter plate of 7 setae.

Limb IV. Pre-epipodite setulated. Epipodite ovoid. Exopodite subquadrangular, with six setae; seta 1–3 long, of similar length; setae 4–6 about two times shorter; seta 5 longer than seta 6. Setae 1–4 flat, plumose; setae 5–6 thin, with short setulae in distal portion. Inner portion of limb IV with four setae and small sensillum (Fig. 1J). Scraping seta (1) slender, sharp; flaming-torch setae (2–4) with well-developed distal portion, bearing 6–7 long setulae each; first flaming-torch seta (2) wider than the two other (2–3). Gnathobase with two-segmented seta and a small hillock distally. Gnathobase filter plate of 5 setae.

Limb V. Preepipodite setulated, epipodite oval. Exopodite oval, with four plumose setae. Setae 1–3 long, subequal in length, seta 4 short, three times shorter than seta 1. Inner lobe as oval lobe. At inner face, two setae, distal seta two times longer than basal. Filter plate V absent.

Ephippial female. Habitus similar to that in parthenogenetic female, with a bit elongated body (body height/ body length = 0.56–0.67), ephippium slightly additionally pigmented in brown, with a single egg.

Male. Body (Fig. 1K, 4A) low oval, with maximum height at the middle, height/length ratio about 0.65. Valve with naked setae at postero-ventral angle, followed by a submarginal row of strong spinules of similar size.

Postabdomen (Figs. 1L, 4B–C) of moderate size, slightly narrowing in anal portion, subrectangular in postanal portion. Length about 3.1 height. Postanal angle not defined, preanal angle well-defined. Postabdominal claw situated on small protrusion in ventral portion of convex distal margin. The sperm ducts open above the protrusion at distal margin of postabdomen. Dorso-distal margin of postabdomen straight; distal part of postabdomen 1.35 times longer than preanal; postanal portion equal or slightly shorter than anal portion. Clusters of short setulae in place of marginal denticles; lateral fascicles of setulae same as in female. Postabdominal claw weakly curved, shorter and more robust than in female. Basal spine about 0.3 length of claw.


FIGURE 3. Alona setulosa arangureni ssp. nov. from Laguna Navio Quebrado, Colombia (type locality), parthenogenetic

female. A, distal portion of mandible. B, IDL of thoracic limb I. C, endites of limb I. D, basal portion of limb I. E, limb II. F,

exopodite of limb III.

Antennule broader than in female, with 10 long terminal aesthetascs and 2 lateral aestetascs of similar length. Male seta subterminal, arising at 1/4 length from tip, not reaching the end of antennule.

Thoracic limb I (Fig. 4D) with U shaped copulatory hook with its free arm little longer than basal one. Row of 14–17 short setulae on ventral face of limb under copulatory brush. First IDL seta absent; 2nd and 3rd setae subequal in length, 2 times shorter and thinner than in female; male seta of same size as 2nd seta.

Size. Length of adult parthenogenetic females 0.32–0.42 mm, of ephippial females 0.32–0.38 mm, of adult males 0.28–0.29 mm.


FIGURE 4. Alona setulosa arangureni ssp. nov. from Laguna Navio Quebrado, Colombia (type locality), adult male. A, lateral

view. B–C, postabdomen. D, copulatory hook of limb I.

TABLE 1. Main morphological differences between O. setulosa arangureni ssp nov., O. setulosa setulosa (Megard,

1967) and O. glabra (Sars, 1901).

Variability. A single parthenogenetic female without connection between posterior and central main head pore was found.

Differential diagnosis. O. setulosa arangureni ssp. nov. differs from O. setulosa setulosa (Megard, 1967) (see Sinev, 2009) by: (1) greater IP/PP ratio in both sexes; (2) posterior margin of labrum without clusters of short setulae; (3) longer postabdomen, with length/height ratio about 2.7–3; (4) male postabdomen with straight dorsal margin in whole distal portion. O. setulosa arangureni ssp nov. is similar to O. glabra (Sars, 1901) in general shape and morphology of postabdomen, but, like nominative subspecies, differs from it by interrupted connections between head pores. For other differences between these taxa see Table 1. Like nominative subspecies, A. setulosa

O. setulosa arangureni ssp. nov. O. setulosa setulosa O. glabra

Female

Body height/body

length

0.66–0.76 0.64–0.67 0.62–0.66

Ventral margin of

valves

with 40–45 setae With 35–45 setae With 40–50 setae

PP/IP ratio 0.4–0.5 IP 0.3–0.4 IP 0.2–0.3

Major head pores Connection interrupted in several

places

Connection interrupted in

several places

Connection continuous

Distance from lateral

head pore to midline

1.1–1.5 IP 1.2 IP 0.9–1 IP

Labral keel Without clusters of setules on the

posterior

margin

With clusters of setules on the

posterior

margin

Without clusters of setules on

the posterior

margin

Length of postanal

portion of postabdomen

About 1–1.1 of anal portion length About 1 of anal portion length About 1.2–1.4 of anal portion

length

Distal angle of

postabdomen

Acute or right Right Acute

Flaming-torch setae of

limb IV

With developed distal portion With developed distal portion With reduced distal portion

Male

Postabdomen Of same height along all anal

portion, anal portion of dorsal

margin parallel to ventral margin

Height weakly increases basally

in anal portion

Height strongly increases

basally in anal portion

Length/height ratio of

postabdomen

2.8–3 2.5 2.6

Marginal setules of

postabdomen

All setules in postanal groups

short, of same length

All setules in postanal groups

short, of same length

Distal setules in postanal

groups long, length of setules

decreases basally


arangureni ssp nov. clearly differs from other South American Ovalona species: O. altiplana (Kotov, Sinev & Berrios, 2010); O. kaingang (Sousa, Elmoor-Loureiro & Santos, 2015); and O. nigra (Smirnov, 1996) by interrupted connection between main head pores.

Distribution and ecology. To the date, the subspecies is known to the date from La Guajira Department in

Colombia. The type locality, the Laguna Navio Quebrado, is a small (surface area of 10.7 km2) lagoon system, with shallow water bodies (depth 0.3–0.7 m) covered by submerged and floating aquatic vegetation; temperature varies over the seasons in the range of 28 and 31 °C; during the sampling, pH was 7.8–8.3 and salinity between 0–28 PSU, being 0 in the mouth of the river and 28 near the communication to the sea.

Discussion

Our study revealed few differences between females and male of the North American and Colombian populations of Ovalona setulosa (see Sinev 2009). The morphology of appendages is the same but morphology of labrum and of male and female postabdomen is quite different. O. setulosa is distributed from North Canada (Chengalath 1982) to North Mexico and Central Mexican Plateau (Elías-Gutiérrez et al. 2008; Sinev & Silva-Briano 2012). However, O. setulosa is absent in the South regions of Mexico (Elías-Gutiérrez et al. 2008). The level of the differences between North American and Colombian populations, in our opinion, is not high enough to justify separation of the latter into separate species, but, together with geographical disjunction, fully justifies placement of Colombian population into a separate subspecies.

In general appearance and morphology of postabdomen, O. setulosa arangureni ssp. nov. is somewhat similar to the most common Neotropical species of the genus, O. glabra. These species can be easily recognized by morphology of main head pores, but can be easily confused if head pores are not studied. However, there are two important characters which clearly show that O. setulosa arangureni ssp. nov. is not a form of O. glabra with aberrant morphology of head pores. Unlike O. setulosa arangureni (and O. setulosa setulosa as well), O. glabra

has: (1) flaming-torch setae of limb IV with reduced distal portion (see Sinev, 2001a) and (2) male postabdomen strongly widening basally in anal portion, with enlarged postanal marginal setules; length of distalmost setule in each group is about width of the base of postabdominal claw (see Sinev 2001a; Sinev & Silva-Briano 2012).

O. setulosa arangureni ssp. nov. is also somewhat similar to two other South American Ovalona species, the endemic of Andes O. altiplana and the recently described Brazilian O. kaingang. However,both these species have uninterrupted connection between pores and differs from O. setulosa arangureni ssp. nov. in morphology of postabdomen. O. altiplana have shorter postabdomen with longer postanal denticles, length of five distalmost ones exceed the width of postabdominal claw base (Kotov et al. 2010). O. kaingang have broadly rounded distal angle of postabdomen and more narrow, widely spaced postanal lateral fascicles of setulae; distance between fascicles about half or more fascicle width (Sousa et al. 2015a).

At the moment, five species of Ovalona are known from South America: O. glabra is distributed through the whole continent, including Andes; O. nigra and O. altiplana are endemics of Andes (Sinev, 2015); O. kaingang is known from South-East Brazil (Sousa et al. 2015a); and O. setulosa arangureni ssp. nov. so far only known from Colombia. Populations of Ovalona from Venezuela described by Rey & Vásquez (1986) as A. karelica and later by Van Damme et al. (2011a) as Alona cf. karelica have questionable status, and can belong to a separate species, but presently should be treated as a form of O. glabra (Sinev 2015). O. weinecki (Studer, 1878) is known from subantarctic islands, including Falklands, but not recorded in continent.

Our results shows that Alona-like Chydoridae in Neotropic are far from fully studied and certainly deserve further investigations. In respect of cladoceran studies, the most intensively investigated region of South America is Brazil south of Amazon basin. Many local faunas of the area were recently investigated in detail (Hollwedel et

al. 2003; Elmoor-Loureiro 2007; Van Damme & Dumont 2010; Debastiani-Junior et al. 2015 and others). Recent studies revealed here a number of new Chydoridae species (Sinev & Elmoor-Loureiro 2010; Elmoor-Loureiro 2014; Sousa et al., 2015ab), and even a new genus (Elmoor-Loureiro et al. 2013). In contrast, modern studies of Colombian Cladocera are just started, but already revealed three new taxa of subfamily Aloninae: Ovalona setulosa

arangureni ssp. nov., Leydigia lourdesae Kotov & Fuentes-Reinés, 2014; and Leberis colombiensis Kotov & Fuentes-Reinés, 2015 (Kotov & Fuentes-Reinés 2014, 2015), all these taxa so far are not recorded in Brazil. On the other hand, a number of other Aloninae species are known from Brazil only (Sinev & Elmoor-Loureiro 2010;


Sousa et al. 2015ab). Many species of South American Aloninae are widely distributed, inhabiting the whole Neotropic province (excluding Andes), or even penetrate into the South of Nearctic region. However, our data suggests that a number of endemics with limited areas of distribution can be present in South America as well, and areas north and south of Amazon basin can have different local species of Aloninae, and of other groups of Cladocera as well.

Acknowledgements

We are very grateful to Dr. Alexei A. Kotov for his valuable consultations. This work is supported by the Russian Science Foundation grant 14-14-00778.

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