ZOOTAXA. Monograph. Magnolia Press Auckland, New Zealand. Accepted by M. Alonso: 15 Mar. 2010; published: 19 Apr. 2010

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1 Zootaxa 2430: 1 66 (2010) Copyright 2010 Magnolia Press Monograph ISSN (print edition) ZOOTAXA ISSN (online edition) ZOOTAXA 2430 The Cladocera (Crustacea: Branchiopoda) of six high altitude water bodies in the North Chilean Andes, with discussion of Andean endemism ALEXEY A. KOTOV 1, ARTEM Y. SINEV 2 & VIVIANA LORENA BERRIOS 3 1. A. N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow , Russia; alexey-a-kotov@yandex.ru 2. Department of Invertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Leninskie Gory, Moscow, , Russia. artem_sinev@ mail.ru 3. Departamento de ciencias del Mar, Universidad Arturo Prat, Casilla 121, Iquique Chile. viviana6771@gmail.com Magnolia Press Auckland, New Zealand Accepted by M. Alonso: 15 Mar. 2010; published: 19 Apr. 2010

2 ALEXEY A. KOTOV, ARTEM Y. SINEV AND VIVIANA LORENA BERRIOS The Cladocera (Crustacea: Branchiopoda) of six high altitude water bodies in the North Chilean Andes, with discussion of Andean endemism (Zootaxa 2430) 66 pp.; 30 cm. 19 Apr ISBN (paperback) ISBN (Online edition) FIRST PUBLISHED IN 2010 BY Magnolia Press P.O. Box Auckland 1346 New Zealand Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN ISSN (Print edition) (Online edition) 2 Zootaxa Magnolia Press KOTOV ET AL.

3 Table of contents Abstract... 3 Abbreviations... 4 Introduction... 4 Material and methods... 5 Results... 6 General... 6 Descriptions of and comments on selected taxa... 8 Daphnia (Ctenodaphnia) paggii sp. nov... 8 Ilyocryptus cf. nevadensis Cervantes-Martínez, Gutiérrez-Aguirre & Elías-Gutiérrez, Ilyocryptus denticulatus denticulatus Delachaux, Macrothrix atahualpa Brehm, Macrothrix cf. oviformis Ekman, Pleuroxus fryeri sp. nov Pleuroxus hardingi Smirnov, Kotov & Coronel, Pleuroxus varidentatus Frey, Alona glabra Sars, Alona nigra Smirnov, Alona altiplana sp. nov Coronatella cf. circumfimbriata (Megard, 1967) Geoffreya gen. nov Geoffreya fryeri gen. nov., sp. nov Leydigia (Leydigia) louisi louisi Jenkin, Discussion High endemism of high altitudes Some relicts in the Andes: possible signatures of the Mesozoic Era Andean-Patagonian relationships Conclusions Acknowledgements References Abstract It was recently demonstrated that mountain areas add to a significant increase of the number of known cladoceran species due to higher chances of endemism. We studied six water bodies in the North Chilean Andes (XV, I and II Regions of Chile), located at more than 4000 m above sea level, and found 19 species of the Cladocera. Several selected taxa are redescribed in detail. Three new species are described: Daphnia (Ctenodaphnia) paggii sp. nov. from Salar de Lagunillas, Pleuroxus fryeri sp. nov. from Crater Lake in Licancabur Volcano (type locality) and Laguna Leija, Alona altiplana sp. nov. from Crater Lake in Licancabur Volcano (type locality) and three other water bodies. A single female, determined as Coronatella cf. circumfimbriata (Megard, 1967) from Salar de Lagunillas, probably also belongs to a yet undescribed species. Geoffreya fryeri gen. nov., sp. nov. is found in a single locality, Salar de Lagunillas. The genus Geoffreya gen. nov. (Chydoridae: Aloninae) is different from all other genera of Aloninae in a set of unique and rare characters, such as (1) exopodites III and IV, both having only four setae, all of them terminal, (2) seta 1 of exopodite V longer than setae 2 4, (3) six setae in filter plate II; (4) small size of exopodites III V in comparison with limb I; (5) two interconnected main head pores of different size; (6) clusters of hard setules on basal and middle segments of antenna II exopodite, (7) very long setules in the lateral fascicles on the postabdomen. The postabdominal claw in Geoffreya gen. nov. bears a very short basal spine. Among 19 species found, four taxa can be regarded as cosmopolitan, ten (more than half among 19!) are Andean endemics (among them six have Patagonian congeners), two are Andean-Patagonian, one circum-neotropical, one Gondwanian, one found in two very distant localities, in the Andes and high mountains of Mexico. Some zoogeographical patterns revealed in this study are discussed. Some species groups, i.e. Daphnia paggii sp. nov. and its conge- CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 3

4 ners, Ilyocryptus nevadensis and its congeners, are probably differentiated from some pan-continental Mesozoic taxa. We agree with previous authors that the Andean-Patagonian patterns in the cladoceran distribution likely has a Last Glacial Maximum (LGM) signature (Löffler 1984; Adamowicz et al. 2002; Mergeay et al. 2008). Based on the marked increase of number of endemic Andean taxa after our investigation of only six water bodies, we assume that the number of Neotropical endemics in the Cladocera could be significantly increased in course of further studies of the Andean high altitude waters. Key words: Crustacea, Anomopoda, fauna, taxonomy, systematics, new species, new genus, biogeography, South America, Neotropics Abbreviations Collections. AAK, personal collection of A. A. Kotov, Moscow, Russia. ALMG, Albany Museum, Grahamstown, U.S.A. AM, Australian Museum, Sydney, Australia. DGF, Collection of D. G. Frey, Support Center of the Smithsonian Institution Museum of Natural History in Suitland, Maryland, U.S.A. ECO-CH-ZOO, Zoological collection of the Chetumal branch of ECOSUR, Chetumal, Q. Roo, Mexico. GOS, Collection of G. O. Sars, Zoological Museum of the Oslo University, Norway. LPM, La Plata Museum, Argentina. MGU, Zoological museum of the Moscow State University, Moscow, Russia. NHM, The Natural History Museum, London, United Kingdom. UNAM, Reference Collection of National Autonomus University of Mexico, Mexico city, Mexico. USNM, The Smithsonian Institution Museum of Natural History, Washington, D.C., U.S.A. In text and illustrations. IDL, inner distal lobe of limb I. IP, interpore distance. ODL, outer distal lobe of limb I. PP, postpore distance. Introduction Problems in cladoceran taxonomy and the poor state of biogeography in the group, are mainly connected with most researchers still ignoring the obvious, so that acceptable decisions about relationships cannot be made without describing and comparing pertinent populations in detail (Frey 1993: P. 186). There are about 620 known species of the Cladocera in the world fauna, but their diversity is significantly underestimated, and level of study is very different for different continents (Forró et al. 2008). Unfortunately, only a limited portion of non-european territories is studied adequately, according to standards of the 21th century, at least with accurately determined species, such as Thailand (Maiphae et al. 2005), Cape Province of South Africa (Smirnov 2008), South Mexico, Belize and Guatemala (Elías-Gutiérrez et al. 2006), Central Mexico (Elías- Gutiérrrez et al. 2008b), Brazilian Pantanal (Hollwedel et al. 2003). Species composition of some other regions remains a mystery for the limnologists, or is preliminary information only for some groups, e.g., for Daphnia. This situation strongly compromises any attempts for further biogeographical analysis (see discussion in Kotov 2008a). For example, Green (1995) analysed the altitudinal distribution of the cladocerans in Africa and South America (especially in Africa, in attempt to compare distribution with latitude), but some of his taxa are complexes of species with different biology and altitudinal-latitudinal preferences, so, new analysis using updated data could significantly change his conclusions. For most regions we have information from publications with accurate determinations (e.g. Smirnov & Timms 1983), but these are outdated and need revision due to a rapid progess of systematics in many cladoceran groups, or data from more recent publications based on morphology or genetics with unknown (and frequently quite poor) quality of the species determination. Recently a program of Barcoding of Life involved the cladocerans from different regions (Elias-Gutierrez et al. 2008a; Quiroz-Vázquez & Elías-Gutiérrez, 2009), but we are still a long way from coordinating such genetical data with an adequate taxonomy, requiring great efforts of both experts in taxonomy and molecular phylogeny, acting according to their own models. It is 4 Zootaxa Magnolia Press KOTOV ET AL.

5 a difficult task to use a phylogenetic tree in taxonomy, but this quite obvious idea is usually ignored by many molecular biologists. Also it is obvious fact that attempts to abandon the benefits of morphological studies in favor of a limited molecular identification system would ultimately impede our understanding of biodiversity (Will & Rubinoff, 2004). High mountains contribute significantly to an increase of the number of known cladoceran species due to higher chances of endemism (Kotov 2008a, b). Several micro-endemisc, known up to recently only from a single water body, were described from mountains of North America (Ciros-Pérez & Elías-Gutiérrez 1997; Cervantes-Martínez et al. 2000), South America (Hann 1986), Africa (Kotov & Štifter 2006; Sinev 2009a) and Asia (Kotov 2008a, b), although we can expect that for some of these taxa new localities will be found in the future, and ideas on their distribution may be changed. Cladocera of South America, especially of Argentina and Chile, are being studied currently by Juan C. Paggi (Paggi 1977, 1993, 1996, 1998, 1999). Some authors also made a contribution to taxonomy (Korovchinsky 2004; Kotov & Gololobova 2005) and molecular phylogeny (Adamowicz et al. 2004, 2009) of several cladoceran groups. The Andes is a very specific part of South America, attracting significant efforts of former and recent authors (Delachaux 1919, 1928; Brehm 1924, 1936; Harding 1955; Villagra de Gamundi 1984, 1986; Valdivia Villar, 1988). Some scarce information on the diversity of the Andean cladocerans can be extracted from two preliminary Chilean checklists (Ruiz & Bahamonde 1989; Berrios & Sielfeld 2000). The aim of this communication is to describe cladocerans found in high altitudes of the Chilean Andes (six water bodies, all above 4000 m above sea level) and to discuss their biogeographic relationships. Material and methods The Salar de Huasco and Salar de Lagunillas (localities 3 4 in Table 1) were collected by VB, quantitative samples were taken from these water bodies using standard nets and standard methods of collection (Dumont & Negrea 2002). These water bodies are located in the Altiplano region of Tarapaca. This area is characterized by mean precipitations of 150 mm/year, mean evaporation rates of 2500 mm/year (López & Garcés 2002) and mean yearly temperatures of 5ºC (Risacher et. al. 1999). Precipitations show a high seasonality, determining wet (rainy) periods during summer and a dry period during autumn, winter and spring. The Lagunillas basin has a surface area of 129 km 2, and the saline lagoon has a surface area of 0.2 km 2. The Huasco basin has a surface area of 1572 km 2 and its saline lagoon has a surface area of 51 km 2. Both basins have shallow water bodies, mainly saline, and lagoons of a non-saline water or aquifers that feed the remaining water bodies. They all show variable size depending on the season of the year. The studied samples were taken from adjacent areas of saline and non-saline athalassic waters. The latter ones arise from bofedales, high altitude pastures with abundant plant species where water drains from the upper springs towards the lower saline lagoons. The faunal composition of the bofedales shows successions of its biological components, which change depending on the optimal salt concentration for each case. The species found at the Salar del Huasco were registered in a permanent lagoon with a mean depth of 20 cm. The concentration of total dissolved solids (TDS) registered in the water column fluctuated from 0.27 g/l and g/l, being classified as non-saline to meso-saline (Hammer et al. 1983). Lower saline concentrations were registered mainly during wet periods (summer), favouring the presence of cladocera and other non saline aquatic invertebrates. In the Lagunillas the species were registered in a temporal shallow lagoon, with a mean depth of 10 cm. However in 2001 and 2002 depths of 20 cm were registered, caused by an exceptional inflow of water coming from spring rains. In the water column the salt concentration of g/l was registered, which does not show seasonal differences. This lagoon is classified as sub-saline to hypo-saline (Hammer et al. 1983). Material on other water bodies was obtained from Dr J. Gibson and Dr F. Fiers, see Table 1 for further information. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 5

6 TABLE 1. LIST OF SAMPLES STUDIED IN THIS PAPER. Region N Locality Coordinates Altitude, m.a.s.l. Collectors Date of collection Reference Décima Quinta Region de Arica y Parinacota 1 Laguna Chungará 18º15 S; 69º10 W 4520 F. Billiet & B. Jadin 06.xi.1991 Pontificia Universidad Católica de Chile (2008); Dorador et al Décima Quinta Región de Arica y Parinacota 2 A small stream in wet prairie, Parinacota Region 18º10 S; 69º15 W 4390 F. Billiet & B. Jadin 06.xi.1991 No information Primera Región de Tarapacá 3 Salar de Lagunillas 19º59 S; 68º54 W 4253 V. L. Berrios 2001 and 2007 Pontificia Universidad Católica de Chile (2008) Primera Región de Tarapacá 4 Salar de Huasco 20º18 S; 68º50 W 4165 V. L. Berrios , different dates Pontificia Universidad Católica de Chile (2008) Segunda Región de Antofagasta 5 Crater Lake, summit of Licancabur Volcano, Atacama Desert 23º29 50 S; 67º41 38 W 5883 V. Gaete 04.ii.2007 Hann (1986) Segunda Región de Antofagasta 6 Laguna Leija 23º30 S; 67º542 W 4325?? Grosjean (1994) For determination, specimens were selected from preserved samples under a binocular stereoscopic microscope, and studied under an optical microscope in a drop of a glycerol-formaldehyde mixture. At least five parthenogenetic females from each locality (if possible) were dissected under a stereoscopic microscope for the study of appendages and postabdomen. Drawings were prepared using a camera lucida attached to an Alphaphot compound microscope. For two new taxa, five females of each species were lyophilised, mounted on an aluminium stub, coated with gold, and examined under a scanning electron microscope (SEM). These specimens were dissected, coated with gold, and examined under SEM again. A system of enumeration for different setae on thoracic limbs proposed by Kotov (2000a, b) for alonine chydorids and then tested for other anomopods (Ishida et al. 2006; Kotov 2007b) is used. Here we use it for the first time with reference to various anomopods from the same region, and think that this system may be acceptable for all cladoceran groups, which seems to be a special methodological result of this paper. Results General Among 19 species found in six water bodies (Table 2), only four taxa can be regarded now as cosmopolitan ones. Each of them apparently consists of a series of species (of unknown number), so, any ideas on their distribution are premature (see Chilean records in Ruiz and Bahamonde 1989). Preliminary molecular-genetical data on Chydorus sphaericus, Daphnia pulex and Simocephalus vetulus species groups immediately revealed a series of cryptic taxa, masking under these names, even in the Holarctic (Belyaeva & Taylor 2009; Elías- Gutiérrez et al. 2008; Adamowicz et al. 2009). See also a history of revision of Ilyocryptus sordidus species group using morphological approach (Kotov & Štifter 2006). With our material, we cannot give an adequate conclusion on identity/conspecifity of our Andean populations with C. sphaericus, D. pulex, S. vetulus and I. sordidus s.str., this is a task for further global revisions. 6 Zootaxa Magnolia Press KOTOV ET AL.

7 Ten taxa found here (more than half among 19!) are Andean endemics; among them six (more than half of ten) have Patagonian relatives. Other revealed zoogeographical patterns are more rare, namely: Andean- Patagonian taxon (2); circum-neotropical taxon (1); Gondwanian taxon (1), and is found in two distant localities, in the Andes and high mountains of Mexico (1). TABLE 2. List of species found in studied samples. Family, subfamily N Taxon Localities Comments on taxon distribution Daphniidae 1 Daphnia (Ctenodaphnia) chilensis Hann, Known only from its type locality, at present seems to be a micro-endemic 2 Daphnia (Ctenodaphnia) paggii sp. nov. 3 Known only from its type locality, at present moment seems to be a micro-endemic, having the closest relative in the southern half of South America (D. menucoensis) 3 Daphnia (Daphnia) cf. pulex Leydig, Simocephalus cf. vetulus (O. F. Müller, 1776) Ilyocryptidae 5 Ilyocryptus cf. nevadensis Cervantes- Martínez, Gutiérrez-Aguirre & Elías- Gutiérrez, Ilyocryptus denticulatus denticulatus Delachaux, Regarded as cosmopolitan, but apparently consists of a series of species 4 Regarded as cosmopolitan, but apparently consists of a series of species 2 Now known from two very distant localities above 4500 m.a.s.l. in Mexico and Chile, but identity must be checked 1, 2 Andean endemic, while other subspecies inhabits Patagonia 7 Ilyocryptus cf. sordidus (Liévin, 1848) 1 Regarded as cosmopolitan, but, probably consists of a series of species Macrothricidae 8 Macrothrix atahualpa Brehm, , 3 Andean endemic having closest relative (M. magna) in Patagonia 9 Macrothrix cf. oviformis Ekman, Andean-Patagonian taxon Chydoridae: Chydorinae 10 Chydorus cf. sphaericus (O. F. Müller, 1776) 4 Regarded as cosmopolitan, but, probably consists of a series of species 11 Pleuroxus freyeri sp. nov. 5, 6 Andean endemic, having a closest relative in Patagonia (P. scopuliferus) 12 Pleuroxus hardingi Smirnov, Kotov & Coronel, , 4 Andean endemic 13 Pleuroxus varidentatus Frey, Andean-Patagonian taxon, possibly has a North American congener Chydoridae: Aloninae 14 Alona glabra Sars, , 4 Neotropical 15 Alona altiplana n. sp. 1, 3, 4, 6 Andean endemic 16 Alona nigra Smirnov, , 4 Andean endemic, closest relatives inhabit North America 17 Coronatella cf. circumfimbriata (Megard, 1967) 3 Possibly a new species, known from a single locality, at this moment seems to be a micro-endemic 18 Geoffreya fryeri gen. nov., sp. nov. 3 Known only from its type locality, at this moment seems to be a micro-endemic 19 Leydigia louisi louisi Jenkin, , 4 Neotropical-Afrotropical ( Gondwanian ), distributed North up to southern Mexico; other subspecies in Valley of Mexico - appearance of the latter seems to be a result of recent colonisation from Neotropics CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 7

8 Description of selected taxa Order Anomopoda Sars, 1865 Family Daphniidae Straus, 1820 Daphnia (Ctenodaphnia) paggii sp. nov. (Figures 1 4) Etymology. This species is dedicated to Dr Juan C. Paggi, renowned Argentinean investigator of the South American Cladocera. Type locality. Salar de Lagunillas (Table 1: locality 3). Holotype. Adult parthenogenetic female, MGU Ml 83. Label of the holotype: "Daphnia paggii sp. nov., 1 parth. from Lagunillas Salt Lake, North Chile, coll. in summer 2002, HOLOTYPE". Allotype. Adult male, MGU Ml 84. Paratypes. Five parthenogenetic females, MGU Ml 85; three males, MGU Ml 86; one ephippial, one preephippial and three parthenogenetic females, AAK All type specimens are in 90% alcohol. Short diagnosis. Parthenogenetic female. Body almost transparent, subovoid, lacking a caudal spine, a distinct depression between head and rest of body. Rostrum short and rounded, ventral margin of head with shallow pre-ocular and post-ocular depressions, eye capsule located significantly below the level of anteriormost point of head; ocellus present Head shield with slightly projected, rounded fornices, a projection from valves penetrates to about 1/2 of head shield length. Secondary fornix short. A group of relatively long setae in middle of ventral margin, short setae at postero-ventral margin of valve, no setules between them. First abdominal segment with a relatively short (as long as postabdominal claw) process, slightly bent anterior; second segment with a small process, the third segment with a very low, mound-like process; the fourth segment lacking a process. Preanal angle of postabdomen ill-defined, postanal angle not expressed. On outer side of postabdominal claw, the first and second (proximal) pectens and the third (medial) pecten consisting of numerous thin teeth, each approximately two times shorter than claw diameter; the fourth pecten consisting of numerous fine setules not reaching the tip of claw. Body of antenna I well-developed, tips of aesthetascs projected beyond tip of rostrum, antennular sensory seta arise from base of mound of the antenna I and not reaching tip mound. On limb I setae 1-3 long, bearing short setules distally, seta 4 long, with long setules. Limb II with seta 1 somewhat longer than half of a soft seta near it; gnathobase with four anterior setae and posterior setae. Limb III with seta 2 of exopodite bearing short setules; inner-distal portion of limb III with setae 1, 2 and 3 long, seta 4 shorter than each of the latter. Gnathobases III IV with numerous posterior soft setae. Limb V with exopodite supplied with two distal setae and a long, curved lateral seta. Ephippium darkly pigmented, "O"-shaped, two eggs with axes almost parallel to main body axis and located at a very acute angle to dorsal margin, anterior process long, but a posterior process fully absent, postero-dorsal portion of valves incorporated into ephippium. Adult male. Body elongated, dorsal margin of valves almost straight, very slightly elevated above head, distinct depression between head and valves, postero-dorsal angle distinct, but without a caudal spine. Head with rounded rostrum. Abdomen without processes on each of its segments. Postabdomen with distal portion as a short tube, gonopore opens subdistally, without a genital papilla. Antenna I long, antennular sensory seta thin, located distally on end of antenna I body; flagellum long, bisegmented, its distal segment setulated.. IDL of limb I with a bent copulatory hook, and two setae of very different size; anterior setae 2, 3 and 4 smaller that these in female and supplied with long, fine setules. On distalmost endite of limb II, anterior seta 1 slightly bent and asymmetrically setulated. Size. Females up to 2.40 mm, males mm 8 Zootaxa Magnolia Press KOTOV ET AL.

9 Description. Adult parthenogenetic female. Body almost transparent, subovoid in lateral view, with maximum height in middle, anteriormost and posteriormost extremities of body lies at the level of longitudinal body axis, dorsal margin of valves elevated above head, regularly convex (Fig. 1A), a distinct depression between head and rest of body. Postero-dorsal angle in adults lacking a caudal spine, as a rounded triangle (while smaller females bear a very short caudal spine), ventral margin regularly convex. In ventral view body ovoid, moderately compressed laterally (Fig. 1B). FIGURE 1. Daphnia paggii sp. nov., adult parthenogenetic female from Salar de Lagunillas, Chile (locality 3). A B. Lateral and ventral view; C D. Head, lateral view; E F. Head, ventral and dorsal view; G. Medium portion of valve ventral margin, inner view; H. Postero-ventral portion of valve; I J. Postabdomen, lateral view; K L. Postabdominal claw. Scale bars 1 mm for A B; 0.1 mm for D L. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 9

10 Head with a short, rounded rostrum (Figs 1A, C D), without a pre-rostral fold (see Ishida et al. 2006; Kotov et al. 2006); posterior margin of head straight; ventral margin of head with shallow pre-ocular and postocular depressions (Fig. 1D, arrows). Eye capsule located significantly below the level of anteriormost point of head. Compound eye relatively small, ocellus distinct, although minute, located far from base of antenna I. In ventral view, head with two depressions, delimiting its topmost portion (Fig. 1E, arrows). Head shield widest in middle, with slightly projected, rounded fornices, a projection from valves penetrates to about 1/2 of its length (Fig. 1F, arrow). Labrum with a wide, fleshy main body supplied distally with series of setules, and a setulated distal labral plate (Fig. 1C, arrow). FIGURE 2. Daphnia paggii sp. nov., female from Salar de Lagunillas, Chile. A B. Antenna I, lateral view; C. antenna I, distal view; D. Antenna II, anterior view; E. Sensory setae at coxal portion of antenna II; F G. Distal segments of antennal exopod and endopod; H. Apical swimming seta of antenna II; I J. Ephippial female, lateral and posterior view; K. Dorsal portion of ephippium; L. Sculpture in central portion of ephippium. Scale bars 1 mm for I J; 0.1 mm for A H, K L. 10 Zootaxa Magnolia Press KOTOV ET AL.

11 Carapace subovoid, secondary fornix short (Fig. 1A, arrow), spinules present on whole dorsal margin and on posterior half of ventral margin. A group of relatively long setae in middle of ventral margin (Fig. 1G), short setae at postero-ventral margin of valve (Fig. 1H), no setules between them as in some other species of Daphnia (see Alonso 1996; Ishida et al. 2006). Abdomen relatively short, consisting of four segments, the first (basal most) abdominal segment with a relatively short (as long as postabdominal claw) process, slightly bent forward; the second segment with a small process, the third segment with a very low, mound-like process, covered by transverse rows of minute setules; the fourth segment without a process, with slightly convex dorsal margin (Figs 1I J, each segment is marked by an arrow). Postabdomen elongated, tapering distally, with ventral margin in general slightly convex, lacking of setules (Figs 1I J). Preanal margin long, almost straight, with series of minute setules. Preanal angle ill-defined, postanal angle not expressed. Numerous small paired spines of subequal size on anal portion. Postabdominal seta somewhat longer than preanal margin, its distal and basal segments of similar length. Postabdominal claw regularly bent, with a pointed tip. On outer side, four successive pectens along the dorsal margin (Fig. 1K L, each pecten is marked by arrow): the first and second (proximal) pectens and the third (medial) pecten consisting of numerous thin teeth, each approximately two times shorter than claw diameter; the fourth pecten consisting of numerous fine setules, somewhat shorter than those in the second pecten, not reaching the tip of claw, inner side of claw with uniform setules. Rows of denticles at ventral margin of the claw. Body of antenna I well-developed, with nine aesthetascs of different length terminally (Figs 2A C), their tips projected beyond tip of rostrum, antennular sensory seta (Fig. 2B C, arrow) fine, arise from base of mound of the antenna I, short, not reaching tip of mound. Antenna II with coxal part possessing two short sensory setae of different length (Figs 2D E). Basal segment elongated, distal sensory seta on its posterior face (not represented in our figures) remarkably longer than the basal segment of exopod, minute distal spine at its anterior face (Fig. 2D, arrow). Antennal branches somewhat longer than basal segment, all with series of minute setules. Spines on apical segments rudimentary (Figs 2F G), spine on the second segment of exopod small (its length about half of diameter of third segment) and thin (Fig. 2D). Antennal formula: setae / Swimming setae subequal in size, each seta with basal and distal segments bilaterally setulated, no chitinous insertion within distal segment (Fig. 2H). Maxilla I as a mound with three seulated setae (Fig. 3A). Limb I without accessory seta; outer distal lobe (Fig. 3B: ODL), with a long seta bilaterally armed distally with short setules, and a short second seta; inner distal lobe (Fig. 3B: IDL), or endite 4, with a single, long anterior seta 1, unilaterally armed distally with short setules. Endite 3 with a long anterior seta 2 and two posterior setae (a b). Endite 2 with a long and thin anterior seta 3, bilaterally armed distally with short setules, and two posterior setae (c d). Endite 1 with a long anterior seta 4, armed with long, fine setules distally, and four posterior setae (e h). Two ejector hooks of different length. Limb II with a subovoid epipodite; distal portion as an elongated lobe bearing a soft, distal seta, and a large, soft, lateral seta. Four endites bearing five setae, among them, a stiff anterior seta (Fig. 3D E: 1) somewhat longer than half of the soft seta on this endite, armed with short setules distally, a rudimentary seta (Fig. 3D: arrow) on endite 2 (the third one from distal end). Gnathobase with two rows of setae: four anterior setae (Fig. 3F: 1 4), the longest seta 2 as long as a 'filter plate' seta) and numerous (16 19) posterior setae of gnathobasic filter plate (a p). Limb III with a large, setulated pre-epipodite, ovoid epipodite and a flat exopodite bearing four distal setae (Fig. 3G: 1 4), among them seta 2 distally with short setules, and two lateral (Fig. 3G: 5 6) setae. Innerdistal portion of limb with four endites: endite 4 with a single, long anterior seta, armed distally with short setules (Fig. 3H: 1) and a posterior (Fig. 3H: a) seta, bearing long setules; endite 3 with a single anterior seta (2) and a single posterior (b) seta, both with long setules; endite 2 with a large anterior seta (3) and two posterior setae (c d); endite 1 with a relatively small anterior seta (4) and four posterior (e h) setae. The rest of the limb CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 11

12 FIGURE 3. Daphnia paggii sp. nov., appendages of adult parthenogenetic female from Salar de Lagunillas, Chile, all in anterior view. A. Maxilla I; B C. Limb I; D F. Limb II, its distal portion and gnathobase; G I. Limb III, its inner-distal portion and filtering seta of gnathobase; J K. Limb IV and its inner-distal portion; L. Limb V. Scale bars 1 mm. 12 Zootaxa Magnolia Press KOTOV ET AL.

13 inner-distal portion as a singular large lobe, modified gnathobase, bearing numerous (about 80) posterior soft setae, each with chitinous insertion within basal portion of distal segment (Fig. 3I), a single, relatively long anterior seta (Fig. 3H: 1) in its distal corner, and two short anterior setae (2 and 3) in middle of the filter comb. Limb IV with a large, setulated pre-epipodite, ovoid epipodite and a wide, flat exopodite, bearing four distal and two lateral setae (Fig. 3J: 1 6). Inner-distal portion of this limb with completely fused endites, distally with two setae of unclear homology (Fig. 3K), the most part of limb inner margin is a gnathobase filter plate consisting of numerous (about 55 60) posterior setae. Limb V with a small, setulated pre-epipodite, large, subovoid epipodite, triangular exopodite supplied with two distal setae, among them, more proximal seta rudimentary, and a large, curved lateral seta (Fig. 3L). Inner limb portion as an ovoid flat lobe, with setulated inner margin and a single, large seta. Ephippial female. In contrast to parthenogenetic female, the dorsal margin of the valves is stronger elevated above head, stronger convex in anterior portion and straight in posterior 2/3 of dorsum (Fig. 2I), dorsal wall of carapace additionally chitinised, forming a dorsal plate, supplied with series of fine spinules (Fig. 2K). In anterior view dorsal portion of body strongly compressed laterally (Fig. 2J). Ephippum O-shaped (terminology of Paggi 1996), the most part of ephippium darkly pigmented and covered with sculpturing of small rounded cells, anterior process long, but a posterior process fully absent, postero-dorsal portion of valves incorporated into ephippium. Two resting eggs, axes of which almost parallel to main body axis and located under a very acute angle to dorsal margin, egg chambers not separated from each other. Adult male. Body elongated, in general subovoid, dorsal margin of valves almost straight, very slightly elevated above head, distinct depression between head and valves, postero-dorsal angle distinct, but without a caudal spine (Fig. 4A). Head with a rounded rostrum, a small depression on posterior margin of head ventrally to antenna I joint (Fig. 4B). Ventral margin slightly concave, anterior most extremity completely occupied with optic vesicle, in some males a shallow supra-ocular depression dorsally to it (Fig. 4A, arrow), another shallow depression between first and second bunches of muscles of antenna II (Fig. 4A, arrow). Eye relatively small for a male of this subgenus, ocellus minute. Valve with anterior margin almost straight, supplied with exactly marginal, relatively short setae; anteroventral angle prominent anteriorly, supplied with specially long setae (Figs. 4C D); whole ventral margin with numerous setae located submarginally on inner face of valve (Figs 4C F). Postero-ventral portion of valve with marginal denticles, and short setae located submarginally on inner face of valve, no setules between these setae (Figs. 4G H). Abdomen without processes on each of its segments (Figs. 4I). Postabdomen with preanal margin shorter than in female, expressed preanal angle, and distal portion as a short tube, gonopore opens subdistally, without a genital papilla (Fig. 4J K, arrow). On anal margin, teeth finer than in female. Postabdominal claw short, on its outer surface a two more basal pectens consisting of long and stout setules, and third pecten consisting of numerous finer and shorter setules (Fig. 4L). Antenna I long, slightly and regularly curved, with series of fine setules (Figs. 4B); antennular sensory seta thin, located distally on end of antenna I body (Figs. 4M, arrow); aesthetascs of different length, largest aesthetasc longer than antenna I maximum diameter. Male seta (flagellum) on top of a low, conical, distal (post-aesthetasc) process. This seta long, bisegmented, its distal segment setulated. Antenna II as in female (Fig. 4N), no additional hairs on any segments of exopod or endopod, which are found in many other species. Limb I: ODL large (Fig. 4O), bearing a rudimentary seta and a very large seta supplied with minute setules distally (Fig. 4P); IDL with a bent copulatory hook, and two setae (1 and 1') of very different size, seta 1 naked; in contrast to female, endite 3 with four setae (additional naked seta of unclear homology marked as 2, a small sensillum near it), setae 2, 3 and 4 shorter than those in female and supplied with long, fine setules. Limb II: distal most endite with seta 1 slightly bent and asymmetrically setulated (Fig. 4Q). Size. Holotype 2.10 mm, parthenogenetic females mm; ephippial female 2.40 mm; allotype 1.81 mm; adult males mm. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 13

14 FIGURE 4. Daphnia paggii sp. nov., adult male from Salar de Lagunillas, Chile. A. Lateral view; B. Head; C. Free margin of valve, inner view; D. Setae at antero-ventral angle of valve; E F. Setae at medium portion of valve ventral margin; G. Setae at postero-ventral portion of valve; H. Setae at posterior margin of valve; I. Postabdomen, lateral view; J K. Its distal portion, lateral and dorsal view; L. Postabdominal claw; M. Antenna I, distal portion; N. Antenna II; O P. Limb I and distal portion of large seta on its outer distal lobe; Q. Anterior seta 1 on inner-distal portion of limb II. Scale bars 1 mm for A, C; 0.1 mm for B, D Q. 14 Zootaxa Magnolia Press KOTOV ET AL.

15 Differential diagnosis. Daphnia paggii sp. nov. belong to a group of D. (Ctenodaphnia) species with rounded fornices (Alonso 1985; Glagolev 1986, 1995). D. paggii sp. nov. differs from D. atkinsoni Baird, 1859, D. bolivari Richard, 1888 and D. triquetra Sars, 1903 in absence of a dorsal extension of the carapace into the head shield that forms a heart-shaped lobe. Our new taxon is close to D. mediterranea Alonso, 1985, D. dolichocephala Sars, 1895 and D. salina Hebert & Finston, 1993 (but the former two have projected, not rounded fornices). But specially, D. paggii sp.nov is close to D. menucoensis Paggi, Paggi (1996) analysed the species discrimination within this group in detail, and we agree with his conclusions on the species discrimination, so we are concentrated on differences of our new taxon from D. menucoensis only. Daphnia paggii sp. nov. differs from the former in: (1) distinct ocellus; (2) short and rounded rostrum; (3) very short abdominal processes of first and second abdominal segments; (4) smaller seta of ODL specially short, less that twice thicker than large seta diameter; (5) apparently shorter seta 1 on inner-distal portion of limb II; (6) seta 4 of limb I in female supplied with long setules; (7) dorsal part of carapace of ephippial female strongly curved in anterior portion; (8) a posterior process of ephippium fully absent (rare character in representatives of the subgenus Ctenodaphnia) and postero-dorsal angle of carapace incorporated into ephippium; (9) male gonopores are not split-like; (10) seta 1 of male IDL rudimentary; (11) seta 1 on inner-distal portion of limb II with longer setules. Just characters of gamogenetic specimens seems to be most valuable for differentiation of this species from its congeners, as it was found for other anomopods (Kim et al. 2006; Kotov et al. 2009). Comments. Daphnia (Ctenodaphnia) paggii sp. nov. could not be associated with any of clades revealed in South America by Adamowicz et al. (2004, 2009) genetically, including their undescribed new species. Distribution. D. paggii sp. nov. is known only from its type locality. Apparently this is an Andean high mountain endemic, preferring salt waters as does its closest relative, D. menucoensis (see Paggi 1996). Family Ilyocryptidae Smirnov, 1976 emend. Smirnov, 1992 Ilyocryptus cf. nevadensis Cervantes-Martínez, Gutiérrez-Aguirre & Elías-Gutiérrez, 2000 (Figures 5 6)? Ilyocryptus nevadensis Cervantes-Martínez, Gutiérrez-Aguirre et Elías-Gutiérrez 2000, p , figs Material studied here. Two females from locality 2. Diagnosis of Chilean specimens. Parthenogenetic female. Body triangular-rounded, high, cervical incision shallow, dorsal margin almost straight, postero-dorsal angle expressed (Fig. 5A). No lateral horns or other conspicous structures on valves (Fig. 5B). Moulting incomplete. Head small (Figs. 5C E), with a distinct prominense where dorsal head pore opens, ocellus minute (Fig. 5C), mandibular articulation narrow (Fig. 5E). Labrum with a projected antero-ventral angle (Fig. 5C) and a pair of lateral projections in middle portion (Fig. 5D). Valves with fine reticulation, relatively large dots within cells well observable (Fig. 5F), setae in anterior bunch longer than the rest (Fig. 5G), while setae at postero-ventral portion not remarkably longer than other setae (Fig. 5A). Each seta in middle of ventral margin with specially long setules (Fig. 5H), each seta on posterior margin with markedly thick basal portion terminating in a single spine-like setule, and with finely setulated distal portion (Fig. 5I). Postabdomen high, height maximal in its middle; anus opens much closer to distal extremity than to base of postabdomen (Fig. 5J). Preanal teeth small, numerous, clustered, spaced regularly and standing approximately at right angles with margin (Fig. 5K), a gap between basalmost cluster and basis of postabdominal setae, as it was previously found in few other species (Kotov & Timms, 1998). Rows of small setules near these teeth in basal part of postabdomen. Paired spines large and numerous, this row continuing along preanal margin up to its middle. Large lateral setae as long or even longer than paired spines (Fig. 5L), proximalmost seta located on preanal margin, not far from anus (Fig. 5J). Postabdominal claws relatively robust, no denticles in their distal or middle portions (Fig. 5M). Two spines on CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 15

16 claw base, basalmost member longest. Setules ventrally on claw basal border long. Postabdominal setae long, densely setulated distally with relatively short hairs (Fig. 5N). Antenna I relatively short, proximal segment with a system of rounded hillocks and a terminal crown of conical projections (Figs. 6A B). Distal segment with 4-5 rows of specially small denticles, distal end with a concentric row of hillocks. Two aesthetascs longer than others, and somewhat longer than half distal segment. Antenna II stout, distal burrowing spine almost reaching distal border of basal segment (Fig. 6C). Antennal branches robust, with well-developed denticles FIGURE 5. Ilyocryptus cf. nevadensis, adult parthenogenetic female from a small stream in wet prairie, Parinacota Region, Chile. A. Lateral view; B. Anterior view; C E. Head, lateral, ventral and dorsal view; F. Reticulation on valve; G I. Setae at antero-ventral, ventral and posterior margin of valve; J K. Postabdomen and armature of its basal portion; L. Large lateral setae; M. Postabdominal claw; N. Postabdominal seta. Scale bars 0.1 mm. 16 Zootaxa Magnolia Press KOTOV ET AL.

17 FIGURE 6. Ilyocryptus cf. nevadensis, appendages of adult parthenogenetic female from a small stream in wet prairie, Parinacota Region, Chile. A B. Antenna I, lateral and anterior view; C. Antenna II, anterior view; D. Its exopod. E F. Apical and lateral swimming seta of antenna II; G. Distal portion of limb I. Scale bars 0.1 mm. (Fig. 6D). Apical swimming setae with distal segments feathered along one side by setules of medium size and by minute setules along other side (Fig. 6E). Proximal and distal lateral swimming setae without hooks on their tips, with distal segments armed asymmetrically and with long setules on basal segments (Fig. 6F). Spine on second segment of exopod relatively long, reaching distal end of third segment (Fig. 6D). Limb I with outer distal lobe bearing single large seta. A long seta, bilaterally armed by short setules distally, near ejector hooks (Fig. 6G). Limb VI with inner margin lined by continuous row of long, fine setules. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 17

18 Size in our sample mm (n=2). Ephippial female, male. Unknown. Differential diagnosis. See Kotov and Štifter (2006). Possible differences from Mexican populations. I. nevadensis is described from Lake La Luna in the crater of the volcano Nevado de Toluca, State of Mexico, Mexico, (19 06'13''N, 99 45'20''W, 4680 m.a.s.l.) (Holotype: female, USNM Paratypes: 2 females, USNM ; 1 female, UNAM CL-1072; 2 females, ECO-CH-ZOO 460). Our two females had (1) somewhat longer lateral setae on postabdomen (approximately as long as paired spines, sometimes even longer); (2) finer denticles on sides of postabdomen basally; (3) smaller denticles on antenna I. Unfortunately, two specimens are not enough to speak about a consistence of such differences. In addition, Mexican populations need to be restudied for accurate comparison with other localities, because some fine details were not adequately described by Cervantes-Martínez et al. (2000), see Kotov and Elías-Gutiérrez (2009). Distribution. Previously found only in two adjacent lakes, La Luna and El Sol, in the crater of the volcano Nevada de Toluca, Mexico. Now we added a very distant locality, also in high mountains. The distributional area seems to be disrupted, or the Chilean population may belong to a separate taxon. Ilyocryptus denticulatus denticulatus Delachaux, 1919 Iliocryptus sordidus var. denticulatus Delachaux, 1919, p , Pl. 1, figs Iliocryptus sordidus subsp. denticulatus Delachaux in Smirnov 1976, p. 49, fig. 15 (after Delachaux). Ilyocryptus denticulatus Delachaux in Kotov 2001, p. 190, figs 9 10 (after Delachaux, 1919). Ilyocryptus denticulatus denticulatus Delachaux in Kotov and Štifter 2005, p , figs 1 46; Kotov and Štifter 2006, p , figs Material examined. Many females from localities 1 2. Diagnosis, redescription. Populations from localities 1 2 were described in detail by Kotov and Štifter (2005, 2006). Comments. Type localities of I. denticulatus denticulatus are Lakes Huaron and Lavandera (about 5140 m.a.s.l.), Peru; type material was apparently lost. Distribution. The subspecies is an endemic of Andean highlands in Peru (Lakes Huaron and Lavandera) and North Chile, other subspecies, I. denticulatus freyi Kotov et Štifter, 2005 inhabits Patagonia, where it is quite common. Family Macrothricidae Norman & Brady, 1867 emend. Smirnov, 1992 Macrothrix atahualpa Brehm, 1936 (Figures 7 9) Macrothrix atahualpa Brehm, 1936, p , figs 5 7. Macrothrix palearis Harding, 1955, p , figs 24 33; Uéno 1967, p , figs 27 31; Petkovski 1973, p. 177, 179; Smirnov 1992b, p. 47, 50 51, figs Echinisca palearis (Harding) in Smirnov 1976, p , figs ; Villagra de Gamundi 1984, p , Pls 1 3. Macrothrix atahualpa (Brehm) in Paggi 1993, p Material studied here. 21 females from localities 1 and Zootaxa Magnolia Press KOTOV ET AL.

19 Diagnosis. Parthenogenetic female. Body subovoid in lateral view, dorsal margin without cervical depression; postero-dorsal angle a rounded triangle, located on level of longitudinal body axis. Head ventral margin slightly convex, without ridges. Ocellus very small. Abdomen with a low, robust dorsal process. Dorsal margin of postabdomen distinctly bilobed; preanal margin long, regularly convex, with short transversal series of setules, they became to be shorter basally and fully absent in basalmost part. Reticulation on sides of postabdomen absent. Postabdominal claw regularly bent dorsally, with pointed tip, an external row of 7 10 thin setules in distal half of claw. Postabdominal seta approximately as long as postabdomen, with relatively long distal segment, bilaterally armed with long setules. Antenna I not widened distally, with a low subapical external angulation; sensory seta externally at distance of about 1.5 antennular diameters (at base) from antenna I joint, about 7 8 transverse rows of robust spinules on its inner face, fine spinules at distal end. Lateral seta on proximal endopod segment of antenna II with a series of robust denticles in middle. Limb I without accessory seta; inner distal lobe with three bisegmented setae of different size, largest seta with fine setules, while two other setae armed with strong spinules. A single ejector hook; a fully setulated maxillar process on limb base. A soft seta near scraper 4 on limb II. Exopodite of limb III with small three distal and sole lateral setae; basal endite with only three anterior setae and three posterior setae. Limb IV with exopodite small, bearing only two distal setae of different size; inner distal limb portion with row of only four long, soft setae. Limb V with large, tri-lobed pre-epipodite. Ephippium with two eggs, very dark, with sculpture as a coarse reticulation; dorsal margin almost straight. Adult male poorly described. It is smaller than female, compound eye large, postabdomen in general similar with that in female. Antenna I with two large sensory setae, additional male seta located on special pedestal located somewhat basally than half of antenna I length; difference in size of longest and shorter aesthetascs greater than in female. Size up to 1.15 mm. Redescription of adult parthenogenetic female. Body subovoid in lateral view, maximum height in the middle, height/length = about 0.6, dorsal margin regularly arched from tip of rostrum to posteriormost point, without a cervical depression, carapace not elevated above dorsal margin of head (Fig. 7A). Postero-dorsal angle a rounded triangle, located on level of longitudinal body axis. No particular structures on valves or head. Body moderately compressed laterally. Head large; in lateral view, its dorsal margin evenly convex, no dome above eye; ventral margin slightly convex, without ridges, no projection at base of labrum (Fig. 7B). A special fold runs from mandibular joint anteriorly, it corresponds to a poorly expressed fornix. Compound eye relatively small, ocellus very small (about 1/3 1/4 of eye diameter), located near tip of rostrum. "Dorsal head pore" (dorsal organ) relatively small, ovoid. Labrum large, triangular, with a amall, setulated distal plate. Valve surface with fine reticulation (Fig. 7C). Dorsal margin without serration, ventral margin denticulated. Marginal setae (Figs. 7D F) variable in length and size in different individuals, but order of their sequence: two smaller setae between each bigger one, characteristic also for many other species (Kotov 1999; Kotov et al. 2004; Kotov 2008b). Usually setae at postero-ventral valve portion longest. The order of setal alternation is not too regular in the posterior portion of the ventral margin. Thorax long, abdomen short, with a low, robust dorsal process (Fig. 7G). Postabdomen with a rectangularrounded distal extremity, and with large "heel" basally. Ventral margin almost straight, no series of setules were found there. Dorsal margin distinctly bilobed; preanal margin long, regularly convex, with short transversal series of setules, they became shorter basally and fully absent in basalmost part (Fig. 7G). On anal margin, groups of robust setules, laterally to them series of finer setules. Reticulation on sides of postabdomen absent. Postabdominal claw regularly bent dorsally, with pointed tip, an external row of 7 10 thin setules in distal half of claw, medial row (immediately on ventral margin as seen laterally) of about 3 4 denticles (Fig. 7I), and inner dorsal row with numerous denticles, two of them more robust than the rest (Fig. 7J). Postabdominal seta approximately as long as postabdomen, with relatively long distal segment, bilaterally armed with long setules (Fig. 7K). CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 19

20 FIGURE 7. Macrothrix atahualpa, adult parthenogenetic female from Laguna Chungará, Chile. A. Lateral view; B. Head, lateral view; C. Valve; D F. Setae at antero-ventral and postero-ventral portion of valve; G H. Postabdomen and its basal portion; I J. Postabdominal claw, outer and inner view; K. Postabdominal seta; L M. Antenna I in external view and its distal end in inner view. Scale bars 0.1 mm. Antenna I regularly bent, not widened distally, with a low subapical external angulation (Fig. 7L); sensory seta externally at distance of about 1.5 antennular diameters (at base) from antenna I joint, about 7 8 trans- 20 Zootaxa Magnolia Press KOTOV ET AL.

21 verse rows of robust spinules on its inner face, fine spinules at distal end (Fig. 7M). Nine terminal aesthetascs, two of them strongly larger than the rest, but not longer than 1/2 of antenna I length, each aesthetasc with two minute claws at apex. Antenna II large (Fig. 7A), coxal region folded, with two small basal sensory setae of slightly different size in middle part (Fig. 8A). Basal segment robust, bearing numerous transverse series of spinules; distal sensory seta long, bisegmented; distal burrowing spine longer than proximal segment of exopod. Antennal branches long (about two times longer than basal segment). Swimming setae /1 1 3, spines / Length of apical swimming setae (Fig. 8B) subequal, each seta marked by individual number in Fig. 8A, armature of each seta illustrated in Figs. 8C L. Lateral seta on proximal endopod segment larger than other setae, with a series of robust denticles in middle (Figs. 8J L). Apical spines relatively short (about half of apical segment length or even shorter), slightly curved. A spine on second segment of exopod longer than half of next segment. Large additional denticles on hind side of segments 2 3 of exopod, they are not homologous to true spines in Macrothrix (Kotov et al. 2004; Kotov 2007b) and some other anomopods (Kotov 2006, 2009a). Figure 8. Macrothrix atahualpa, antenna II of adult parthenogenetic female from Laguna Chungará, Chile. A. General view; B. Distal segment of exopod; C I. Apical and lateral swimming setae; J L. Different portions of largest seta of antenna II, located on proximal segment of endopod. Scale bars 0.1 mm. Limb I large, without accessory seta; outer distal lobe (Fig. 9A: ODL) cylindrical, bearing a long apical seta and a short lateral seta. Inner distal lobe massive, with three successive marginal series of setules plus medial series of setules, and three bisegmented setae of different size, largest seta with fine setules, while two other setae armed with strong spinules (Figs. 9B C). Endite 3 posteriorly with slightly curved, setulated seta CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 21

22 (Fig. 9D: a), and straight setae b c; anteriorly on this endite a short, setulated, bisegmented seta 1 (Fig. 9E). Endite 2 with three long bisegmented setae of subequal size (Fig. 9D: d f), each with distal segment bearing fine setules distally and forked seta 2 anteriorly (Fig. 9F). Endite 1 with two bisegmented setae (Fig. 9D: g h) and a fork-like seta 3 (Fig. 9G) anteriorly. A sole ejector hook; a fully setulated seta at inner side of limb base, so-called maxillar process, remainder of gnathobase I (Kotov 1999). FIGURE 9. Macrothrix atahualpa, thoracic limbs of adult parthenogenetic female from Laguna Chungará, Chile. A C. Distal portion of limb II and two smallest setae of its inner distal lobe; D. Limb I, anterior view; E G. Stiff setae on inner portion of limb I; H I. Limb II and distal armature of its gnathobase, posterior view; J K. Limb III and its inner portion, posterior view; L. Limb IV, posterior view; M. Limb V, posterior view. Scale bars 0.1 mm. 22 Zootaxa Magnolia Press KOTOV ET AL.

23 Limb II triangular; epipodite subglobular, exopodite a subovoid lobe with three rows of small setules and short seta distally (Fig. 9H). At inner margin of limb, eight robust scrapers (Fig. 9H: 1 8), scrapers 1 2 with delicate feathering, 3 8 with robust denticles, and small sensillum near scraper 1. Posteriorly to scrapers, a system of low hillocks, and a soft seta near scraper 4. Distal gnathobase with four setae (Fig, 9I: 1 4); filter with four long setae. Limb III with small, globular epipodite; exopodite large and flat, with three distal setae (Figure 9J: 1 3), seta 1 with short setules; setae 2 3 with long setules; lateral group consists of a single seta 4, similar in armature with 2 and 3. Distal endite (see discussion of its homology in Kotov 1999) anteriorly with three bisegmented setae (Fig. 9K: 1 3), unilaterally armed in distal part, small sensillum near each base of seta 1, 2 and 3. Posteriorly, three setae: seta a thick, strong, with robust spinules distally, setae b and c soft, with fine setules distally. Basal endite smaller than distal one. Anteriorly, a bottle-shaped sensillum and only three (Fig. 9K: 4 6) bilaterally setulated setae; posteriorly, only three soft, setulated setae (d f) subequal in size. Gnathobase clearly demarcated from basal endite, with large, bottle-shaped sensillum near border with basal endite (Fig. 9K: 1), and three projections distally (2 4), filter plate fully absent. Limb IV with pre-epipodite small, epipodite very large, globular; exopodite small, with distal group of two bilaterally feathered setae of different size (Fig. 9L). Inner margin of limb with 4 setae (Fig. 9L: 1 4), seta 1 naked, setae 2 4 with inflated basal and elongated distal part, the latter pointed at tip, supplied with relatively robust setules. A small sensillum near base of seta 3. Posteriorly, a row of only four long, soft setae, similar in size, bilaterally setulated from base to tip (Fig. 9L: a d). Distal armature of gnathobase with 4 elements (Fig. 9L: 1 4): a bottle-shaped sensillum (1) near border with basal endite; a large setae 2 with inflated basal segment and elongated, setulated distal segment; a heavy hook 3; and a small, naked seta 4. Posteriorly on gnathobase, a single small seta continues the posterior row of setae of the inner limb face, the sole remainder of filter plate IV. Limb V with pre-epipodite relatively large, tri-lobed (not illustrated in Fig. 9M); epipodite large, globular. A small lobe with single seta remains of the exopodite. Inner-distal portion a large flap, fringed by fine setules, on inner margin three setae, in size increasing distally (Fig. 9L: 1 3). Size up to 1.15 mm. Ephippial female, male. Absent in our material. See Harding (1955) and Villagra de Gamundi (1984), although their descriptions lack many important details. Differential diagnosis. M. atahualpa is a relative of the welll-defined M. paulensis species group (Kotov et al. 2005), sharing with the latter a series of basic synapomorphies (unique or very rare for the genus), first of all (1) large, triangular labrum; (2) a sole ejector hook on limb I; (3) only three anterior and three posterior setae on basal endite of limb III; (4) only two setae on exopodite IV; (5) only four posterior setae on inner portion of limb IV. It differs from members of M. paulensis-group in (1) only somewhat convex ventral margin of head and (2) presence of groups of setules instead of strong spines on inner margin of antenna I. M. atahualpa differs from a relatively similar Australo-African taxon, M. capensis, in a long distal segment of postabdominal seta as it was marked by Harding (1955) and Smirnov (1976). There is a small chance that M. atahualpa is a junior synonym of Patagonian M. magna Daday, 1902, which was earlier regarded as a member of hirsuticornis-group, but the latter idea is now considered incorrect, see Kotov (2007b). At least, M. atahualpa and M. magna are very similar, especially in male morphology (compare illustrations of Daday 1902 and Harding 1955), which is regarded to be specially important for species determination in Cladocera (Kotov 1999; Kim et al. 2006; Kotov 2008b). Unfortunately, the first description of M. magna was lacking many important characters. Our preliminary analysis of Daday s (1902) illustrations led to conclusion that M. atahualpa differs from M. magna in (1) large basal heel of postabdomen; (2) denticles on postabdominal claw located closely and occupying less then half of the claw length; (3) absence of abdominal processes; (4) longer sensory setae on male antenna I. These differences probably indicate existence of two separate taxa (Kotov 1999; 2008b). But our preliminary ideas must be checked with con- CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 23

24 sideration of Daday s types and new samples from Patagonia. In any case, the Andean M. atahualpa has a closest congener in Patagonia. Comments. Type locality of M. atahualpa is unknown water body in the vicinities of Lake Titicaca (Peru- Bolivia border), no further details in Brehm (1936). Brehm did not specify types of his taxa. Some samples of V. Brehm are found and will be deposited to NHM (V. Kořínek, pers. comm), but no information on Titicaca samples there. Both M. atahualpa Brehm, 1936 and M. palearis Harding, 1955 were described from the same area (Lake Titicaca region), and both descriptions were relatively detailed. Probably Harding (1955) did not know Brehm s (1936) paper, which was not cited by him. Without any doubts the Harding s taxon is a junior synonym of Brehm s species. Smirnov (1992b) marked just M. palearis as valid, but listed M. atahualpa in the list of synonyms. Since that time a new edition of ICZN (2000) appeared, and according to several cases of ICZN (2000), M. atahualpa Brehm, 1936 has priority and this name must be used as valid, while M. palearis Harding, 1955 is a junior synonym of the former. Distribution. M. atahualpa is very common in many high mountain water bodies of Peru, Bolivia, North Argentina (Brehm 1936; Harding 1955; Uéno 1967; Villagra de Gamundi 1984; Valdivia Villar 1988; Paggi 1993) and North Chile. Macrothrix cf. oviformis Ekman, 1900? Macrothrix oviformis Ekman, 1900, p , Pl. 4, figs Material examined here. Seven females from locality 3. Comments. Type locality of M. oviformis is... einer Lagune in der Nähe von Rio Turbio (Ekman 1900) in Santa Cruz, Argentina. Type material is, most probably, lost, at least, absent from Ekman s Collection in SMNH (Kotov & Gololobova 2005). We did not find any differences of our Andean population from locality 3 from those of M. oviformis described from southernmost portion of South America (Kotov 2007b). But this conclusion must be checked basing on better material from several localities. Distribution. According to present-day data, M. oviformis seems to be distributed both in Patagonia and the Andes. Family Chydoridae Dybowski et Grochowski, 1894 Subfamily Chydorinae Dybowski et Grochowski, 1894 emend. Frey, 1967 Pleuroxus fryeri sp. nov. (Figures 10 14)? Pleuroxus aduncus (Jurine) in Harding 1955, figs Etymology. This species is named after Dr Geoffrey Fryer, FRS, famous British investigator of the Chydoridae, as well as other Cladocera, other Branchiopoda, other Crustacea and their predators, fish. Type locality. Crater Lake, Licancabur Volcano (locality 5). Holotype. Adult parthenogenetic female in 90% alcohol, MGU Ml 81. Label of the holotype: "Pleuroxus fryeri sp. nov., 1 parth. from Crater Lake, Licancabur Volcano, Chile-Bolivia border, coll. in 04.ii.2007 by V. Gaete, HOLOTYPE". Paratypes. 40 parthenogenetic females, MGU Ml 82; 40 parthenogenetic females, AAK Other material studied. Five parthenogenetic female from locality 6, AAK Zootaxa Magnolia Press KOTOV ET AL.

25 FIGURE 10. Pleuroxus fryeri sp. nov., adult parthenogenetic female from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A B. Lateral and ventral view; C. Head, lateral view; D. Head shield; E. Tip of rostrum; F G. Dorsal head pores; H J. Labrum, labral keel and its apex; K. Antenna I; L. Antenna II; M. Apical seta of antenna II. Scale bars 0.1 mm. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 25

26 Diagnosis. Parthenogenetic female. Body brown, relatively opaque, high for the genus, postero-ventral angle without a tooth. In anterior view, body moderately compressed laterally, with triangular-rounded dorsum, because top of carapace ridged. Rostrum long, protruding downward and slightly posterior. Head shield elongated, its posteriormost portion rounded, postpore distance (PP) = about 3 4 interpore distance (IP). Labrum with a large medial keel having a well-defined apex, protruding behind tip of rostrum. Valves with a distinct reticulation, within polygons a fine punctuation. All setae on ventral margin exactly marginal except of anterior most portion. A row of small setules on posterior valve margin, with bases located exactly marginally. Postabdomen wide, preanal margin slightly concave, shorter than anal margin, the latter concave, preanal and postanal angle expressed, postanal margin approximately as long as anal margin, dorso-distal angle widely rounded and slightly prominent distally. Postanal margin provided with thin postanal teeth, organised in series of 2 4 members. Postabdominal claw massive, with two basal spines, proximal one being half size of distal one or even shorter. Antenna I small, with a well-defined basal peg. Antennular sensory seta arising at one third of antennular length from distal end directly from antenna I body, without any prominences. Antennal formula, setae 0 0 3/1 1 3, spines 1 0 1/ All apical "swimming" setae, as well as basal and distal lateral setae of endopod subequal in size, with chitinous insertions within distal segments. Spine on proximal segment of exopod small. Limb I with accessory seta, outer distal lobe with two setae of different length, inner distal lobe with relatively short, naked first seta, second and third setae subequal in size and similarly armed distally with short setules. Two ejector hooks of subequal size. A remnant of maxillar process with a single seta. Number of setae in filter plates II V 8 9 6(7) 4. Size up to 1.16 mm. Ephippial female, male. Unknown. Description. Parthenogenetic female. Body brownish, relatively opaque. In lateral view body oval, high for the genus (body height/ body length = body height/length = in juveniles, in adults), maximum height in middle (Figs. 10A, 11A). Dorsal margin evenly arched from tip of rostrum to postero-dorsal angle, which is ill-defined, posterior margin slightly convex, postero-ventral angle broadly rounded, without any teeth, ventral margin with a slight prominence in middle. Body moderately compressed laterally (Fig. 11B), with triangular-rounded dorsum, because top of carapace ridged (Figs. 10B, 11C). Head with a long rostrum, protruding downward and posterior (Fig. 10A, C, 11D F). Ocellus smaller than compound eye, lies closer to base of antenna I than to compound eye. Head shield elongated, with maximum width on level of mandibular articulation, its posteriormost portion rounded (Figs. 10D, 11G), rostrum with a terminal tubercle (Figs. 10E, 11F). Two major head pores (Figs. 10F), sometimes a special inner organ associated with head pores is visible under opical microscope and SEM (Fig. 10G, 11H); postpore distance = about 3 4 interpore distance (Fig. 10D). Lateral head pores minute, normally located asymmetrically to midline (Figs. 10F G, 11H). Labrum with fleshy main body, small distal labral plate and a large medial labral keel, with well-defined apex (Fig. 10H), projecting beyong tip of rostrum (Figs. 10C, 11D E) and supplied with 1 2 terminal hillocks (Figs. 10H J). Valves large, with a distinct reticulation (Fig. 12A), within polygons a fine punctuation (Fig. 12B); in dorsal portion lines of reticulation inflated, forming a sculpture as a series of folds (Figs. 13A B), while in ventral portion reticulation also expressed, but not forming such sculpture (Figs. 13C D). Ventral margin armed with numerous setae of different size in different regions, in anterior portion they are naked and located far from margin (Fig. 12C), following setae plumose and located exactly marginally, each of more anterior seta has an expanded basis bearing a spine, following setae longer and lacking expanded bases and spines (Figs. 12D G). A row of minute setules on posterior valve margin, with bases located exactly marginally (Figs. 12G, 13D). Postabdomen wide (Figs. 12H I, 13E), its ventral margin almost straight, preanal margin slightly concave, shorter than anal margin, the latter concave, preanal and postanal angle expressed, postanal margin approximately as long as anal margin, dorso-distal angle widely rounded and slightly prominent distally, inflated basis of claws bordered from postanal margin by a distinct depression (Figs. 12J, 13F). Postanal margin provided with thin postanal teeth, organised in series of 2 4 members, they are larger in its distal portion. 26 Zootaxa Magnolia Press KOTOV ET AL.

27 FIGURE 11. Pleuroxus fryeri sp. nov., adult parthenogenetic female from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A. Lateral view; B. Ventral view; C. Anterior view; D. Lateral view of female with removed valve; E F. Head, lateral and ventro-lateral view; G H. Head pores. Scale bars 0.1 mm for A G, 0.01 mm for H. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 27

28 FIGURE 12. Pleuroxus fryeri sp. nov., adult parthenogenetic female from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A B. Valve and its surface; C G. Setae at different portions of valve margin; H I. Postabdomen; J L. Postabdominal claw. Scale bars 0.1 mm. 28 Zootaxa Magnolia Press KOTOV ET AL.

29 Laterally on postabdomen series of short, fine setules. Postabdominal seta somewhat longer than preanal margin, with distal segment bearing short, rare setules (Fig. 12I). Postabdominal claw shorter than anal margin, massive, evenly curved, with setules along ventral margin, and two basal spines, proximal one being half size of distal one or even shorter (Figs. 12J L, 13F). Antenna I small, length about 2.5 less than labral keel length, narrowing distally (Figs. 10C, K, 13G), with a well-defined basal peg. Antennular sensory seta slender, approximately as long as half the antenna I, arising at one third of antennular length from distal end directly from antenna I body, without any prominences. Nine short aestetascs of slightly differing size. Antenna II (Figs. 10L, 13H) relatively short, coxal part with two sensory setae, basal segment robust, with a rudimentary distal spine. Antennal branches relatively elongated, endopod longer than exopod, all segments cylindrical, antennal formula, setae 0 0 3/1 1 3, spines 1 0 1/ All apical "swimming" setae, as well as basal and distal lateral seta of endopod subequal in size, with chitinous insertions within distal segments (Fig. 10M). Spine on proximal segment of exopod small. Exopod and endopod apical spines subequal in size. Limb I (Figs. 13H, 14A D). Distal portion with accessory seta, outer distal lobe (ODL) bears a long terminal seta with distal segment unilaterally armed with minute setules, and a short lateral seta with bilaterally setulated distal segment. Inner distal lobe (IDL) with relatively short, naked first seta, second and third setae subequal in size and similarly armed distally with short setules. Endite 3 with three posterior setae (a c) and a setulated anterior seta 1 (Fig. 14B), all of subequal length. Endite 2 with three posterior setae d f, and thin anterior seta 2 bilaterally armed with short setules distally (Fig. 14C), a small sensillum near the latter. Endite 1 with thin posterior setae g i, a short seta j, and anterior seta 3 (Fig. 14D) somewhat shorter than seta 2, with a sensillum neat its base. Fascicles of thin setules on inner face of limb, plus bunches of longer, thicker setules at ventral margin of limb. Two slender ejector hooks of subequal size. A remnant of maxillar process with a single seta. Limb II (Figs. 14E H). Exopodite small, subquadrangular, with a short seta. Inner portion of limb with eight scrapers (Fig. 14E: 1 8), 1 2 longer and armed with fine setules (Fig. 14G), others armed distally with thin spinules (Fig. 14H), setae 6 8 shortest. A system of small projections posteriorly to scrapers 2 4, and a small sensillum near scraper 4. Distal armature of gnathobase with a bunch of setules and four setae (Fig. 14F), a bunch of stout setules distally to seta 1, a bunch of fine setules basally to it. Filter plate II with eight setae. Limb III (Fig. 14I). Epipodite ovoid, exopodite sub-rectangular, with four distal setae (1 4) and three lateral setae (5 7). Distal endite with three anterior setae (Fig. 14J: 1 3), basalmost seta (3) shorter, small sensillae near bases of setae 2 and 3. Basal endite with four anterior setae (Fig. 14J: 4 7), slightly increasing in size basally, armed with fine setules distally, a small bottle-shaped sensillum near seta 4. On posterior surface, six soft setae (Fig. 14I), bilaterally armed with fine setules. Gnathobase not clearly separated from basal endite. Distal armature of gnathobase with a large, bottle-shaped sensillum, three setae, and a bunch of setules (Fig. 14J: 1 4), filter plate III with nine setae. Limb IV (Fig. 14K L). Pre-epipodite rounded, setulated; epipodite without a finger-like projection; exopodite wide, subovoid, with seven setae of unequal size (Fig. 14K: 1 7). Inner-distal portion of limb IV with four marginal setae (Fig. 14L: 1 4). Distalmost seta (1) stout, naked, setae 2 4 setulated, with thick distal segments, sensillae located near setae 2 and 3. On posterior surface, four soft setae (Fig. 14K). Distal armature of gnathobase with four setae (Fig. 14L: 1 4), filter plate with six, rarely seven setae. Limb V (Fig. 14M). Pre-epipodite setulated; epipodite subovoid, without a finger-like projection; exopodite large, subovoid, with a single distal seta 1 and three lateral setae (2 4), distally to seta 1 there are two projections bearing long setules. Inner limb portion as elongated, flat lobe, with setulated inner margin, supplied with setulated setae 1 and 2. Distal armature of gnathobase as a single projection, filter plate V with four long setae. One to two parthenogenetic eggs in brood pouch. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 29

30 FIGURE 13. Pleuroxus fryeri sp. nov., adult parthenogenetic female from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A B. Sculpture of valve in its dorsal portion; C D. Valve, ventral portion; E F. Postabdomen and postabdominal claw; G. Antenna I and antenna II; H. Limb I, outer view. Scale bars 0.1 mm. 30 Zootaxa Magnolia Press KOTOV ET AL.

31 FIGURE 14. Pleuroxus fryeri sp. nov., adult parthenogenetic female from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A. Limb I; B D. anterior setae on its endites; E. Limb II; F. Distal armature of its gnathobase; G H. Scrappers 1 and 4 of limb II; I. Limb III; J. Its inner portion; K. Limb IV; L. Its inner portion; M. Limb V. Scale bars 0.1 mm. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 31

32 Ephippial female, male. Unknown. Size. Holotype 1.13 mm, parthenogenetic females mm. Differential diagnosis. Among aduncus-like taxa of the southern hemisphere (Frey 1993; Smirnov et al. 2006), only P. fryeri sp. nov. and P. scopuliferus Ekman, 1900 have (1) top of carapace ridged (triangular in anterior view); (2) filter plate of gnathobase III of nine setae. The former differs from the latter in (1) head shield rounded posteriorly; (2) rostrum shorter, not projecting behind apex of labral keel; (3) no denticles at postero-ventral angle; (4) quite projected peg on antenna I; (5) significantly greater size. P. fryeri sp. nov. is the fourth species of South American Pleuroxus which is brown, not transparent. It differs from P. scopuliferus as described below, from P. paraplesius Frey, 1993 having no denticles at posteroventral angle, from P. hardingi Smirnov et al., 2006 in well-developed basal peg of antenna I and all setae on ventral margin exactly marginal. Distribution. It is known only from two localities, Crater Lake in Licancabur Volcano, and Laguna Leja, both in Atacama Desert, North Chile. Harding (1955) illustrated "P. aduncus" from Lagunillas pond in vicinities of Lake Titicaca, habitually similar to our taxon. Unfortunately, description and figures are lacking many important details, and we can finally confirm the presence of P. fryeri sp. nov. in the vicinities of Lake Titicaca. In any case, P. fryeri sp. nov. is an Andean endemic. Pleuroxus hardingi Smirnov, Kotov & Coronel, 2006 Pleuroxus hardingi Smirnov, Kotov & Coronel, 2006, p , figs Material examined here. Many females from localities 3 and 4. Diagnosis, description, differential diagnosis. See Smirnov et al. (2006). Comments. Type localities are pools of the bofedal system in the cordillera del Tunari (part of the Cordillera Oriental) near the city of Cochabamba, Cercado Province, Bolivia ( m.a.s.l., 17 10'56"S 17 17'19"S, 66 07'62" 66 22'99"W). Holotype: female, MGU Ml 57. Paratypes: 5 females, MGU Ml 58. Distribution. Andean endemic, common in highland water bodies. It was known from pools in the cordillera del Tunari (Bolivia), and from Lagunillas Pond near Lake Titicaca (Bolivia-Peru border) (Smirnov et al. 2006). Here we found two populations from North Chile, so, this taxon is wider distributed in Andes than previously assumed. Pleuroxus varidentatus Frey, 1993 Pleuroxus varidentatus Frey, 1993, p , figs ; Smirnov 1996a, p , figs ; Smirnov et al. 2006, p , figs Material examined here. Five females from locality 4. Diagnosis, description, differential diagnosis. See Frey (1993), Smirnov et al. (2006). Comments. Type locality of P. varidentatus is a marshy channel alongside a small wooden church in Pta. Bandera, Santa Cruz Province, Argentina" (Frey 1993). Holotype. female, LPM. Paratypes. 2 females, ALMG GEN 951A, 5 females, ALMG GEN 951B; 2 females, AM P40973; 5 females, AM P40974; 2 females, GOS 21292a, 5 females, GOS 21292b; 2 females, NHM , 5 females, NHM , 5 females, NHM ; 2 females, USNM ; 5 females, USNM Distribution. It was described from the Province of Santa Cruz, South Argentina (Frey 1993), and then found in the Province of Jujuy, in the northern part of the Argentinean Andes (Smirnov et al. 2006). Now we add a North Chilean record, also from Andean highlands. Habitually similar specimens were found in Mexi- 32 Zootaxa Magnolia Press KOTOV ET AL.

33 can tropical lowlands (Elías-Gutiérrez et al. 2006, 2008b), but these populations are strongly isolated from the Andean those and need to be accurately revised. Subfamily Aloninae Dybowski et Grochowski, 1894 emend. Frey, 1967 Alona glabra Sars, 1901 Alona cambouei Guerne & Richard in Richard 1897, p , figs 35 36; Delachaux 1919, p. 28, Pl. 2, fig. 10; Harding 1955, p , figs ; Uéno 1967, p. 559, figs Alona glabra Sars, 1901, p , Pl. 9: figs. 6, 6a; Daday 1905, p , Pl. 11, figs 3 4; Albertina Kameya 1986, p , fig. 6 7; Sinev 2001a, p , figs 1 40; Elmoor-Loureiro et al. 2004, p. 417, figs Alona pulchella King in Infante 1980, p , figs 5a c. Studied material. 6 parthenogenetic females from locality 3; 75 parthenogenetic females from locality 4. Diagnosis, redescription. See Sinev (2001a). Comments. Type locality of A. glabra is "Argentina", no further information was given by Sars (1901). Lectotype. female, GOS F12326a; Paralectotypes. 40 females, F12326b. Morphology of the studied specimens completely agrees with the recent redescription of the species (Sinev, 2001a) with one exception. About one third of the studied specimens had tuberculated head shields and dorsal part of valves. Polymorphism in the sculpture of head shield and valves is known for several species of Alona-like animals: Alona verrucosa Sars, 1901; A. guttata Sars, 1862; A. rustica Scott, 1895; Coronatella rectangula (Sars, 1862), and others. Apparently specimens with tuberculated valves and head shield should be treated not as a subspecies, but as varieties (Sinev 2001b). Distribution. Common in Neotropics (Sinev 2001a; Elías-Gutiérrez et al. 2006). Alona nigra Smirnov, 1996 (Figures 15 18) Alona nigra Smirnov, 1996b, p , figs Material studied here. 32 parthenogenetic females, 4 ephippial females, 1 adult male, 1 juvenile male of instar II from locality 3; 24 parthenogenetic females from locality 4. Emended diagnosis. Parthenogenetic female. Body regular ovoid, of moderate height, in adults height/ length ratio about 0.7, maximum height at the middle; body moderately compressed laterally. Dorsal margin convex, postero-dorsal and postero-ventral angles broadly rounded. Postero-dorsal angle with about 100 setules organised into groups of setules in each. Ventral margin with about 55 setae. Head shield with broadly rounded posterior margin, rostrum short and rounded. Three disconnected major head pores. PP = IP. Minute lateral head pores located about 1.5 IP distance from midline, at level of middle major head pore. Labrum of moderate size, labral keel of moderate width, with convex anterior margin and a rounded apex. Pre-ultimate abdominal segment with five-six transverse rows of long setules. Postabdomen of moderate width, with parallel margins in postanal portion, length about height. Dorsal margin almost straight in postanal portion, anal margin concave; distal part of postabdomen two times longer than preanal one, with postanal portion about 1.5 times longer than anal one. Postanal portion of distal margin almost straight, anal portion weakly concave. Preanal angle well-defined, postanal angle not defined, distal margin convex, dorso-distal angle broadly rounded. Postanal margin with well-developed marginal denticles, frequently with one-two spinules on anterior margin, near them lateral fascicles of setules, distalmost seta of each fascicle largest, in distalmost fascicles 1.5 times longer than a marginal denticles. Postabdominal claw slightly longer than preanal portion of postabdomen. Basal spine about 0.2 of length of claw. Antenna I CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 33

34 short and wide, with nine terminal aestetascs. Antennal formula setae 0 0 3/1 1 3, spines 1 0 1/ Seta arising from basal segment of endopod longer than endopod. Spine on basal segment of exopod longer than middle segment. Apical spines 1.5 times longer than apical segments. Limb I with accessory seta 1.5 times shorter than ODL seta. IDL with three setae, seta 1 well-developed, about 1/3 length of seta 3. Exopodite of limb III with seven setae, seta 5 being longest. Exopodite IV with six setae. Exopodite V with four setae, filter plate V absent. Epipodites IV and V without projections. Limb VI absent. Male. Body low oval, height/length ratio about 0.64, maximum height at the middle. Body moderately compressed laterally. Dorsal margin convex, postero-dorsal and postero-ventral angles broadly rounded. Postabdomen short and wide, narrowing distally, dorso-distal angle not defined. Preanal angle not defined, postanal angle weakly-defined. Distal part of postabdomens 1.8 times longer than preanal one. Gonopore openings located almost at the end of postabdomen. Wide clusters of short setules in place of marginal denticles. Postabdominal claw very short, 2.5 times shorter than preanal portion of postabdomen, with blunt apex bearing small spine, basal spine long, sinuous, about 1/3 of claw length. Antenna I with ten terminal and two lateral aestetascs, male seta arising at 1/4 length from tip, about 1/4 of antenna I length. Limb I with V-shaped copulatory hook. IDL seta 1 two times smaller than in female, setae 2 and 3 subequal in length, three times shorter and thinner than in female, male seta large, hook-like, 1.5 times longer than seta 3. Size up to 0.65 mm in females and up to 0.42 mm in males. Redescription. Parthenogenetic female. In lateral view body regularly ovoid, of moderate height, maximum height at middle of body (Figs 15A B). In adults height/ length ratio about 0.7, in juveniles about 0.63 in of instar I, about 0.65 in instar II (Figs. 15C D). Dorsal margin uniformly curved, postero-dorsal and postero-ventral angles broadly rounded, posterior margin uniformly curved, ventral margin in general straight (Fig. 15E). Antero-ventral angle rounded. Both head and valves without any sculpture. Head of moderate size, triangle-round in lateral view, rostrum short, pointing downward. Eye and ocellus large, of subequal size. Distance from tip of rostrum to ocellus similar with that between ocellus and eye. Head shield with maximum width behind mandibular articulation, without any prominent sculpture (Fig. 16A). Rostrum short, more narrow than in most other species of the genus, broadly rounded. Posterior margin of head shield broadly rounded, slightly wavy. Three disconnected major head pores (Figs 16A B), comparative size of pores variable. PP about IP. Lateral head pores minute, located about 1.5 IP distance from midline, at the level of middle major head pores. Labrum of moderate size. Distal labral plate without setulation. Labral keel wide (heigh less than 1.5 width), with rounded apex (Figs 16C E). Anterior margin of keel convex, slightly undulated, posterior margin without any setules. Valve subovoid. Ventral margin almost straight, with about 55 setae, first 15 setae long, about ten next setae short, other setae of moderate length (Fig. 15E). A row of about hundred setules along the posterior margin (Fig. 15F), at some distance from one on inner side of carapace, these setules not organized into groups. Postero-dorsal angle bears about hundred short setules organised in groups of setules in each, distalmost seta in each group being longest and thickest. Thorax and abdomen subequal in length, dorsal surface of abdominal segments not saddle-shaped. No abdominal projections. Penultimate abdominal segment with five-six transverse rows of long setules (Fig. 15G). Postabdomen of moderate width, with parallel margins in postanal portion, convex distal margin and broadly rounded dorso-distal angle, length about height (Figs 15H I). Ventral margin almost straight. Inflated basis of claws bordered from distal margin by clear incision. Dorsal margin with distal part two times longer than preanal one, with postanal portion about 1.5 times longer than anal one. Postanal portion of distal margin almost straight, anal portion weakly concave. Preanal angle well-defined, postanal angle not defined. Preanal margin weakly convex. Postabdomen with well-developed marginal denticles, frequently with one-two spindles near base on anterior margin (Fig. 15J), and with three-five groups of marginal setules on 34 Zootaxa Magnolia Press KOTOV ET AL.

35 FIGURE 15. Alona nigra, parthenogenetic female from Salar de Lagunillas, Chile. A B. Large adult; C D. Juvenile; E. Armature of valve free margin; F. Posterior margin; G. Abdomen; H I. Postabdomen; J. dorso-distal angle of postabdomen; K. Postabdominal claw. Scale bars 0.1 mm. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 35

36 anal margin and eleven-twelve lateral fascicles of setules on anal and postanal margin, distalmost seta of each fascicle largest, in postanal portion, longest setule in each fascicles 1.5 times longer than a marginal denticles. Additional row of three-five fascicles between lateral fascicles and marginal setules in preanal region. Postabdominal claw slightly longer than preanal portion of postabdomen. Basal spine about 0.2 of length of claw (Fig. 15K), 2-3 setules basally to it on postabdominal claw. Antenna I relatively short and wide, not reaching tip of rostrum, with three clusters of short setules at anterior face (Fig. 16F). Antennular sensory seta slender, two times shorter than antenna I, arising at 2/3 distance from the base. Nine terminal aesthetascs, two longest of them about 1/2 length of antenna I. All aesthetascs projecting beyond anterior margin of head shield. FIGURE 16. Alona nigra, parthenogenetic female from Salar de Lagunillas, Chile. A. Head shield; B. Dorsal head pores; C E. Labrum; F. Antenna I; G H. Antenna II. Scale bars 0.1 mm. Antenna II short and massive (Figs. 16G H). Antennal formula, setae 0 0 3/1 1 3, spines 1 0 1/ Basal segment robust, branches short and broad, basal segments of both branches 1.5 times longer than others. One of terminal setae shorther than two others. Seta arising from proximal segment of endopod longer than endopod. Seta arising from middle segment of endopod of similar size with apical setae. Spine on basal segment of exopod longer than middle segment. Apical spine 1.5 times longer than apical segments. 36 Zootaxa Magnolia Press KOTOV ET AL.

37 Thoracic limbs. Five pairs. Limb I of moderate size (Figs. 17A B). Epipodite was not found, accessory seta of moderate length. ODL (Fig. 17A), with a long seta, armed with minute setules in distal part; IDL with three setae and four-five clusters of small setules on ventral face, setae 2 and 3 with subequal in length, similar in length to ODL seta, both with thin setules in distal part, seta 1 of about 1/3 length of ODL seta. Endite 3 with three posterior setae (a c), and anterior seta 1 shorter than others. On endite 2 there are three setae (d f), setae e f robust, armed with robust setules in distal part, seta d shorter than ODL setae, and similar to endite 3 setae. Endite 1 with two 2- segmented setae (g h), both setulated in distal part, without a flat seta shifted to limb base, present in some other species. A small senssillum on anterior face of limb on both endites 1 and 2. Five rows of thin long setules on ventral face of limb. Two ejector hooks, one of them somewhat larger than the other. Maxillar process with a short seta. Limb II (Figs. 17C) with exopodite elongated, supplied with a short, naked seta (Fig. 17D). Eight scraping setae (1 8), increasing in length distally, scrapers 3 and 5 armed with more robust setules than others. Distal armature of gnathobase with four setae. Filter plate with seven setae, the distalmost one considerably shorter. Limb III with exopodite trapezium-shaped, with seven setae (Fig. 17E: 1 7). Seta 5 being longest, setae 1 and 2 about 1/3 and 1/4 length of seta 5, respectively, other setae shorter, seta 4 geniculated. Setae 1 and 2 armed with short setules in distal part. Distal endite with three setae (Fig. 17E: 1 3), two distalmost members slender, sharp, with distal parts unilaterally armed with sharp denticles; basalmost seta 3 much shorter, bilaterally armed with long setules. Basal endite marginally with four stiff setae (Fig. 17F: 4-7), increasing in size in basal direction. Four soft posterior setae (a d) increasing in size basally, a small sensillum near the base of distalmost seta. Gnathobase not clearly separated from basal endite. Distal armature of gnathobase with four elements: the first one elongated, cylindrical sensillum, second thin, geniculate seta, others two sharp spines. Filter plate III with seven setae. Limb IV with exopodite rounded, supplied with six plumose setae (Fig. 17G: 1 6). Seta 4 longest, setae 5 6 and 2 of about 2/3 length of seta 4, setae 3 and 1 of about 1/3 length of seta 4. Inner-distal portion of limb IV with four marginal setae (Fig. 17G: 1 4), distalmost seta slender, sharp, armed with sharp denticles, three flaming-torch setae of similar shape, armed with long setules, decreasing in size basally. Three soft setae (Fig. 17G: a c) increasing in size basally. Gnathobase with a large sensillum, a 2-segmented seta, and pair of small hillocks distally. Filter plate with five setae. Limb V (Fig. 17H) with setulated pre-epipodite, epipodite was not found; exopodite oval, not divided into two lobes, with four plumose setae, decreasing in size basally, seta 1 four times shorter than seta 4. Inner limb portion an oval lobe, with setulated inner margin. At inner face, two setae (1 2), 1 two times longer than 2. Filter plate absent. Ephippial female (Fig. 18A). Body subrectangular, dorsal and posterior margins of valves almost straight, postero-dorsal angle expressed, ephippium dark brown, almost black. Male. Body shape of instar II juvenile male (Fig. 18B) of same shape as juvenile female of same instar. Adult male is very small in comparison with the adult female, with lower body (Fig. 18G). General shape low oval, body height/body length about Both eye and ocellus significantly larger than in female, ocellus larger than eye. Postabdomen of instar II juvenile male more narrow, than that of female, with subrectangular distal portion (Fig. 18C). Gonopore openings located close to the end of postabdomen. Marginal denticles same as in female. Postabdominal claw 1.5 times shorter than preanal portion of postabdomen, basal spine same as in female. In adult male postabdomen short and wide, narrowing distally, dorso-distal angle not defined (Fig. 18H). Preanal angle not defined, postanal angle weakly-defined. Distal part of postabdomen 1.8 times longer than preanal. Gonopore openings located almost at the end of postabdomen. Wide clusters of short setules in place of marginal denticles, in five distalmost clusters setules much thicker than in others. Lateral fascicles of setules similar to these of female, but 1.5 times narrower. Postabdominal claw very short, 2.5 times shorter CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 37

38 than preanal portion of postabdomen, with blunt apex bearing characteristic small spine, basal spine long, slightly sinuous, about 1/3 of claw length (Fig. 18I). FIGURE 17. Alona nigra, thoracic appendages of parthenogenetic female from Salar de Lagunillas, Chile. A B. Limb I, outer (posterior) and inner (anterior) view; C D. Limb II and its distal portion; E F. Limb III and its basal endite; G. Limb IV; H. Limb V. Scale bars 0.1 mm. 38 Zootaxa Magnolia Press KOTOV ET AL.

39 FIGURE 18. Alona nigra, gamogenetic specimens from Salar de Lagunillas, Chile. A. Ephippial female; B F. Juvenile male II, its postabdomen, antenna I; distal portion of limb I and copulatory brush on limb I; G L. adult male, its postabdomen, postabdominal claw, antenna I, limb I and its distal portion. Scale bars 0.1 mm. Antenna I of instar II juvenile male broader than in female, with 9 terminal aesthetascs and an anlage of male seta (Fig. 18D). In adult male, antenna I of same length, but broader than in female, with 10 terminal aesthetascs of similar length and two long lateral aestetascs, male seta arising at 1/4 length from tip, about 1/4 of antenna I length (Fig. 18J). Limb I of instar II juvenile male with U-shaped copulatory hook (Fig. 18E), ventral face of limb with an anlage of copulatory brush seta and a peculiar hillock above it, not present in adult male (Fig. 18F). IDL with anlage of male seta, seta 1 same as in female, seta 2 and 3 much shorter than in female. In adult male, copulatory hook V-shaped, its free arm (= terminal part) little longer than basal one (Figs. 18K L). A group of five long setules located under copulatory brush, at some distance of it a row of about short, robust setules on ventral face of limb. IDL seta 1 two times smaller than in female, setae 2 and 3 subequal in length, three times shorter and thinner than in female, male seta large, hook-like, 1.5 times longer than seta 3. Size. Juvenile females of instar I mm (height mm); juvenile females of instar II mm (height mm); adult females mm (height mm). Single studied juvenile male of instar II 0.42 mm (height 0.27 mm), single studied adult male 0.42 mm (height 0.26 mm). CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 39

40 Differential diagnosis. Alona nigra can be differentiated from the majority of other species of Alona s. lat. belonging to a small set of species with disconnected major head pores. There are only three other known species with this peculiarity (1) paleotropical A. cambouei Guerne & Richard, 1893, (2) Mediterranean A. nuragica Margaritora, 1971, and (3) Australian A. setuloides Smirnov & Timms, No other species with disconnected head pores are known from America. A. cambouei differs from A. nigra in (1) smaller size (less than 0.5 mm), (2) narrow postabdomen with parallel margins, (3) prominent, sharp distal angle, and (4) smaller size of the antennal spines (see Sinev 2001c). A. nuragica differs from A. nigra in (1) tapering postabdomen with more numerous denticles and lateral fascicles and (2) lateral pores as small round cavities (Alonso 1996). A. setuloides differs from A. nigra in (1) lower body (height/length ratio about 0.5), (2) longer postabdomen with obtuse distal angle and straight distal margin, and (3) the spine on basal segment of exopodite of antenna II being shorter than middle segment (Smirnov & Timms 1983). Also, Alona nigra differs from these species in a densely setulated penultimate abdominal segment. The male of A. nigra clearly differs from males of these species in (1) shape of postabdomen and (2) very small postabdominal claw with the spine at the apex. Comments. Type locality of this species is Lake Uru-Uru, Oruro Department, Bolivia (3686 m.a.s.l.). Holotype: female MGU Paratypes: females MGU 3571, 35 73, 3574, 3575, 3581, 3587, 3589, 3594, Our specimens completely fit the initial description of the species (Smirnov 1996b). Complete description of appendages was not provided in the initial description. A. nigra shares numerous affinities with the A. pulchella-group, phylogenetic position of which, according to Van Damme & Dumont (2008a) is uncertain, intermediate between Hexalona- and Cornatellabranches. These similarities include: (1) morphology of IDL setae, (2) absence of third seta on endite 1 of limb I, (3) exopodite IV with plumose setae 1 2, (4) absence of filter plate V and (5) absence of limb VI. Male antenna of A. nigra bears two lateral aesthetascs, which are not present in any groups of Hexalona. The aforementioned A. cambouei is also a member of this group. On the other hand, species of the pulchella-group have more narrow and long postabdomen with parallel margins and prominent distal angle, posteroventral margin of valves with setules not differentiated into groups, small accessory seta of limb I, and more broad rostrum. The morphology of male postabdomen of A. nigra also significantly differs from that of pulchella-group. There are two characters rare for Alona s. lat disconnected major head pores and densely setulated middle abdominal segment. There are only three species with such morphology of head pores (see above), and only one, A. setosocaudata Vasiljeva & Smirnov, 1969, with densely setulated abdomen (see Sinev & Kotov 2001). In our opinion, pulchella-group and related species, like A.nigra and A. setulosa (see Sinev, 2009b) belong to the Hexalona-branch of Alona-like animals, having two most important features of Hexalona seven setae on exopodite III and well-developed first IDL seta. Common features between pulchella-group and Coronatella-branch (see Van Damme & Dumont, 2008a) are mostly reductions of limb structures, which independently appears in several unrelated groups of Aloninae (see Kotov, 2000b). Distribution. So far, Alona nigra was found only in saline lakes of the Andes (Bolivia and North Chile); this species seems to be an endemic of the Andes. Alona altiplana sp. nov. (Figures 19 22) Alona pulchella King in Megard 1967, p. 46, figs 29 32; Alona pulchella var. cambouei Guerne & Richard in Dominguez 1973, p. 3 7, figs Type locality. Crater Lake, Licancabur Volcano (locality 5). 40 Zootaxa Magnolia Press KOTOV ET AL.

41 Holotype. Adult parthenogenetic female in 90% alcohol, MGU Ml 95. Label of the holotype: "Alona altiplana sp. nov., 1 parth. from Crater Lake, Licancabur Volcano, Chile-Bolivia border, coll. in 04.ii.2007 by V. Gaete, HOLOTYPE". Paratypes. 20 parthenogenetic females, MGU Ml 96; 40 parthenogenetic females, AAK M Other material studied. 4 parthenogenetic females from locality 4; 13 parthenogenetic females from locality 3; 5 parthenogenetic females from locality 6. Diagnosis. Parthenogenetic female. Body regular oval, of moderately high, in adults height/length ratio about 2/3, maximum height at the middle; body moderately compressed laterally. Dorsal margin convex, postero-dorsal and postero-ventral angles broadly rounded. Postero-dorsal angle with setules not organised into groups. Ventral margin with about setae. Head shield with broadly rounded posterior margin, rostrum short and rounded. Three major head pores with a narrow connection between them. PP less then 0.3 IP. Minute lateral head pores located about 1.2 IP distance from midline, at between anterior and middle major head pore. Labrum of moderate size, labral keel narrow, with convex anterior margin and a rounded apex. Postabdomen of moderate length and width, slightly narrowing distally, length about 2.5 height. Dorsal margin weakly convex in postanal portion, anal margin concave; distal part of postabdomen about 1.5 times longer than preanal one, postanal and anal margins approximately of similar length. Preanal angle well expressed, postanal angle not defined, distal margin straight, dorso-distal angle rounded. Postanal margin with 5 6 well-developed, sharp denticles, some of them with additional denticles near base; near them 5 7 broad lateral fascicles setules, the posteriormost seta of each fascicle longest, of same length as a marginal denticle. Postabdominal claw slightly longer than preanal portion of postabdomen. Basal spine straight or weakly curved, about 0.25 of the claw length. Antenna I of moderate size, with nine terminal aestetascs. Antennal formula: setae 0 0 3/1 1 3, spines 1 0 1/ Seta arising from basal segment of endopod thin, as long as endopod. Spine on basal segment of exopod slightly shorter than middle segment. Spines on apical segments longer than apical segments. Limb I with accessory seta four times shorter than ODL seta. IDL with three setae, seta 1 well-developed, about 1/3 length of seta 3. Exopodite of limb III with seven setae, seta 5 being longest. Exopodite IV with six setae. Exopodite V with four setae, filter plate V absent. Epipodites IV and V without projections. Limb VI absent. Small-sized species, length of adult mm. Description. Parthenogenetic female. In lateral view body regular oval, of moderate height, maximum height at middle of body (Figs. 19A E, 20A), in adults height/length ratio about 2/3. Dorsal margin uniformly curved; postero-dorsal and postero-ventral angles broadly rounded; posterior margin uniformly curved; ventral margin almost straight; antero-ventral angle rounded. Body moderately compressed laterally (Figs. 20B D). Head of moderate size, triangle-round in lateral view, rostrum short, pointing downward (Fig. 20E). Eye larger than ocellus. Distance from tip of rostrum to ocellus in adults slightly greater than that between ocellus and eye, or equal to the latter. Head shield with maximum width behind mandibular articulation, without a reticulation or sculpture; rostrum short, broadly rounded; posterior margin of head shield broadly rounded (Fig. 19F). Three connected major head pores, connection between pores narrow; PP less than 0.3 IP (Figs. 19G H, 20F). Lateral head pores minute, located about 1.2 IP distance from midline, at the level of middle major head pore. Labrum of moderate size; distal labral plate without setulation; labral keel narrow (heigh about than 2.5 width), with rounded apex; anterior margin of keel convex, posterior margin without any setules (Fig. 19I). Valve ovoid, with obscure longitudinal lines, well-visible under SEM (Figs. 20A E, G). Ventral with about setae, first 7 11 setae long, next 10 setae short, other setae of moderate length (Fig. 19J). A row of about 100 setules along the posterior margin, this row located on inner side of valve, setules not organized into groups (Figs. 19K L). Postero-dorsal angle bears short setules of similar length, not organised in groups. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 41

42 FIGURE 19. Alona altiplana sp. nov., parthenogenetic female from Crater Lake, Licancabur Volcano, Chile-Bolivia border (A, L) and Salar de Lagunillas, Chile (B-K). A B. Large adult; C. Juvenile of instar I; D. Juvenile II. E. Minimal size of reproduction; F. Head shield; G H. Dorsal head pores; I. Labrum; J. Free margin of valve; K L. Posterior margin. Scale bars 0.1 mm. 42 Zootaxa Magnolia Press KOTOV ET AL.

43 FIGURE 20. Alona altiplana sp. nov., parthenogenetic female from Crater Lake, Licancabur Volcano, Chile-Bolivia border. A D. Large adult in lateral, ventral, anterior and dorsal view; E. Head; F. Dorsal head pores; G. Sculpture of valves; H. Abdomen. Scale bars 0.1 mm for A E, 0.01 mm for F H. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 43

44 Thorax and abdomen subequal in length, dorsal surface of abdominal segments not saddle-shaped. No abdominal projections. Second abdominal segment with two transverse rows of long thin setules and groups of shorter setules (Fig. 20H). Postabdomen of moderate width, with almost parallel margins in postanal portion, straight distal margin and almost right dorso-distal angle, length about 2.5 height (Figs 19M O, 21A C). Ventral margin straight. Inflated basis of claws bordered from distal margin by clear incision. Dorsal margin with distal part times longer than preanal one, with postanal and anal portions subequal in length. Postanal portion of distal margin straight, anal portion weakly concave. Preanal angle well-defined, postanal angle not defined. Preanal margin weakly convex to slightly sigmoid. Postabdomen with five-six well-developed marginal denticles, frequently with one-two additional minute spinules near base of each denticle, and with four-five groups of marginal setules on anal margin. Eight-eleven lateral fascicles of setules, six-seven distalmost fascicles large, with distalmost seta of each fascicle as long as marginal denticles. Additional row of three-five fascicles between lateral fascicles and marginal setules in preanal region. Postabdominal claw of moderate length, slightly longer than preanal portion of postabdomen. Basal spine about 0.25 of length of claw (Figs 21B, D). Antenna I relatively short and wide, length about 2.5 widht, with two clusters of short setules at anterior face (Figs 21E F, 22A). Antennular sensory seta slender, three times shorter than antenna I, arising at 2/3 distance from the base. Nine terminal aesthetascs, two longest of them about 1/2 length of antenna I. All aesthetascs projecting beyond anterior margin of head shield (Fig. 21E). Antenna II short and massive (Figs 21E, 22B). Antennal formula, setae 0 0 3/1 1 3, spines 1 0 1/ Basal segment robust, branches short and broad, basal segments of both branches 1.5 times longer than others. One of terminal setae shorther than two others. Seta arising from basal segment of endopod of same length with endopod. Seta arising from middle segment of endopod of similar size with apical setae. Spine on basal segment of exopod shorter than middle segment. Apical spines longer than apical segments. Mandibles with asymmetrical molar surfaces as shown in Figs. 21G H. Five pairs of thoracic limbs. Limb I (Fig. 22C), of moderate size; epipodite ovoid; accessory seta very short. ODL (Fig. 22D) with a long seta, armed with minute setules in distal part. IDL (Fig. 22C) with three setae and four-five clusters of small setules on ventral face, setae 2 and 3 2-segmented, shorter than ODL seta, both with thin setules in distal part, seta 1 of about 1/3 length of seta 1. Endite 3 with three posterior setae (a c), and a shorter anterior seta 1. On endite 2 there posterior setae (d f), setae e f long, armed with robust setules in distal part, seta d similar to endite 3 setae a c. Endite 1 with two 2-segmented setae g h, both setulated in distal part, without a flat seta shifted to limb base. Six rows of thin long setules on ventral face of limb. Two ejector hooks, one of them somewhat larger than the other. Maxillar process with a short seta. Limb II with exopodite elongated, supplied with a single naked seta of about 2/3 length of exopodite (Fig. 22E). Eight scraping setae (1 8), armed with spinules of similar shape, increasing in length distally. Distal armature of gnathobase with four elements. Filter plate with seven setae, the posteriormost three times shorter than others. Limb III with oval epipodite; exopodite subrectangular, with seven setae (Fig. 22F:1 7). Seta 5 being longest, seta 2 and 2/5 length of seta 5, other setae short. Seta 2 armed with stronger setules in distal part, seta 1 almost naked, all other setae plumose. Distal endite with three setae (Fig. 22F: 1 3), two distalmost members slender, sharp, with distal parts unilaterally armed with sharp denticles; basalmost seta much shorter, bilaterally armed with long setules. Basal endite with four stiff setae, increasing in size toward the base (Fig. 22G: 4 7), a small sensillum near the base of distalmost seta. Four soft setae increasing in size basally (a d). Gnathobase not clearly separated from basal endite. Distal armature of gnathobase with four elements: an elongated, cylindrical sensillum; thin, bent seta; others two sharp spines. Filter plate III with seven setae. 44 Zootaxa Magnolia Press KOTOV ET AL.

45 FIGURE 21. Alona altiplana sp. nov., parthenogenetic female from from Crater Lake, Licancabur Volcano, Chile- Bolivia border. A C. Postabdomen; D. Postabdominal claw; E F. Head; G H. Molar surfaces of mandibles (right mandible on the right side of photo). Scale bars 0.01 mm. CLADOCERA OF NORTH CHILEAN ANDES Zootaxa Magnolia Press 45