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ZOOTAXA

Revision of Ilyocryptus Sars, 1862 (Cladocera: Ilyocryptidae) of Brazil with description of two new subspecies ALEXEY A. KOTOV1 & LOURDES M. A. ELMOOR-LOUREIRO2 1

A. N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071, Russia. E-mail: [email protected]

2

Laboratorio de Zoologia, Universidade Catolica de Brasilia, QS 7 lote 1, sala M-331, 71966-700, Taguatinga, DF, Brasil. E-mail: [email protected]

Abstract Analyzing numerous samples from Brazil, we found only three previously described species of Ilyocryptus (Cladocera: Anomopoda: Ilyocryptidae): I. paranaensis paranaensis Paggi, 1989, I. sarsi Stingelin, 1913 and I. spinifer Herrick, 1882. Two new taxa, being Neotropical subspecies of the species previously described from Palaearctic, I. cuneatus brasiliensis subsp. nov. and I. silvaeducensis paraensis subsp. nov., are described. Maps of Brazilian records are provided for each species using both original and literature data. Most part of records of Brazilian ilyocryptids concerns I. spinifer, which is among the most common cladocerans in the Neotropics, while other taxa occur significantly rare. I. spinifer has a wide distributional range. “I. sarsi” seems to be a group of close topical congeners, quite possible that I. sarsi s. str. is an exclusively Neotropical taxon. Three other recorded species are Neotropical endemics. We believe that previous records of I. sordidus (Lévin, 1848) from Brazil were misidentifications, probably of two newly described taxa, or I. sarsi. Key words: Branchiopoda, Anomopoda, taxonomy, fauna, South America

Introduction The first record of the Ilyocryptus Sars, 1862 (Cladocera: Ilyocryptidae) from Brazil was made by Ihering (1895), who described Acanthocercus immundus F. Mueller, regarded recently as a junior synonym of I. spinifer Herrick, 1882, which was described from North America (Kotov & Dumont 2000). Since then, the main contributions to study of this genus in Brazil and closest countries were made by Sars (1901), Daday (1905), and Stingelin (1913). Subsequently, the interest to ilyocryptid and generally cladoceran taxonomy was lost, only some records or very scarce descriptions were published (Brandorff et al. 1982). Only in the new millennium, some detailed redescriptions of Brazilian species appeared. A very characteristic Neotropical species, I. sarsi Stingelin, 1913, was redescribed by Kotov et al. (2002a), and its neotype was selected from Sars' sample from Ipiranga, State of Sno Paulo. Many populations of I. spinifer from Brazil were examined by Kotov and Dumont (2000). Subsequently Kotov and Štifter (2004) found a new Neotopical member of the spinifer-group, I. plumosus, but it is not recorded from Brazil. Many authors provided information on new records of ilyocryptids from this country (Brehm 1937, 1938; Schubart 1938, 1942; Bergamin 1939; Green 1972; Hardy 1980; Robertson 1980; Montú & Gloeden 1986; Bohrer et al. 1988; Smirnov & Santos-Silva 1995; Sousa-Dabés 1995; Campos et al. 1996; Nunes et al. 1996; Lansac-Tôha et al. 1997, 1999; Rocha & Por 1998; Rocha et al. 1998; Branco et al. 2000; Crispim & Watanabe 2000; Starling 2000; Espíndola et al. 2000; Morini & Pinto-Silva 2000; Rossa et al. 2001; Sampaio et al. 2002; Azevedo & Bonecker 2003; Hollwedel et al. 2003; Lima et al. 2003; Neves et al. 2003; Serafim Jr. et al. 2003; Elmoor-Loureiro Accepted by M. Alonso: 7 Nov. 2008; published: 12 Dec. 2008

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2004, 2007; Elmoor-Loureiro et al. 2004; Eskinazi-Sant' Anna et al. 2005; Paranaguá et al. 2005; Takahashi et al. 2005; Sousa & Elmoor-Loureiro 2008). The undescribed Neotropical subspecies of I. silvaeducensis Romijn, 1919 was found in the State of Maranhão (Kotov & Štifter 2005a, 2006). Unfortunately, the sole specimen known for that time was dissected and destroyed, so it was not possible to provide a holotype and establish a new subspecies at that time. But recently we found more specimens of this taxon, as well as two populations of a new subspecies of I. cuneatus Štifter, 1988, earlier described from Europe and subsequently found in North America (Kotov et al. 2002b). The aim of this paper is to summarize knowledge of Brazilian Ilyocryptus and describe two new taxa. In each case, our new Neotropical taxon is a closest relative of one, earlier described from Palaearctic. Keeping in mind these especially closest relationships and separate areas of distribution, we prefer to assign a subspecies status to each South American taxon, following a practice of Kotov & Štifter (2005a, 2006). Many recent taxonomists do not recognize a subspecies level, preferring to describe a new species even in case of minute differences. But we, establishing new subspecies, underline some phylogenetic relationships. Apparently this our activity does not contradict with the Code of Zoological Nomenclature.

Methods All operations with ilyocryptids are described in detail by Kotov and Štifter (2006). Maps of distribution were drawn for each species using both original and available literature data from sources listed in the Introduction chapter. Abbreviations for collections. AAK: Personal collection of A. A. Kotov, Moscow, Russia; EL: Personal collection of L. M. A. Elmoor-Loureiro, Universidade Católica de Brasília, DF, Brasil; GOS: collection of G. O. Sars, Zoological Museum of the Oslo University, Norway; MGU: Zoological Museum of the Moscow State University ("Moskovskiy Gosudarstvenniy Universitet"), Russia; MNRJ: Museu Nacional do Rio de Janeiro, Universidade Federal do Rio de Janeiro, Brazil; MZUSP: Museu de Zoologia da Universidade de São Paulo, Brazil.

Results Ilyocryptus cuneatus brasiliensis subsp. nov. Figures 1–3 Etymology. This subspecies is named after the country where it was found, Brazil. Type locality. Itaici - lagoa do pesqueiro (23°06'48.2"S, 47°11'03.0"W), Municipio de Indaiatuba, São Paulo, Brazil. It is a small lake bordered by Eichhornia. The type series was collected on 14.xi.2002 by L.M.A. Elmoor-Loureiro. Type material. Holotype. An adult parthenogenetic female, MZUSP18854. The label of holotype is: "Ilyocryptus cuneatus brasiliensis subsp. nov., 1 parth. & from Itaici - lagoa do pesqueiro, Indaiatuba, SP, Brazil, HOLOTYPE". Paratypes. 5 parthenogenetic females, MZUSP 18855; 5 parthenogenetic females, MNRJ 21506; 5 parthenogenetic females, EL01603; 5 parthenogenetic females, MGU Ml 75; 5 parthenogenetic females, AAK 2008-053. Other material studied. A parthenogenetic female, from Ponte Nova reservoir (appr. 23°34'S, 45°58'W), Município de Biritiba Mirim, São Paulo, Brazil, collected on 02.xii.1997, by M.J.C. Botelho. Short description. Adult parthenogenetic female. General: In lateral view body subovoid, of medium height for the genus (body height/body length = 0.76–0.81 in adults), maximum height in posterior half (Figure 1A–B). Dorsal margin slightly convex to almost straight, postero-dorsal angle distinct. In anterior view,

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KOTOV & ELMOOR-LOUREIRO

FIGURE 1. Ilyocryptus cuneatus brasiliensis subsp. nov., parthenogenetic female from Itaici, Municipio de Indaiatuba, São Paulo, Brazil: A–B—adult female, lateral view; C—anterior view; D—head, lateral view; E—head shield, dorsal view; F—head, ventral view; G—labrum, ventral view; H—free margin of valve; I—antero-ventral margin; J–K—setae at posterior margin. Scale bars 0.1 mm.

REVISION OF ILYOCRYPTUS OF BRAZIL

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FIGURE 2. Ilyocryptus cuneatus brasiliensis subsp. nov., parthenogenetic female from Itaici, Municipio de Indaiatuba, São Paulo, Brazil: A–C—postabdomen, lateral view; D—region of anus; E—anus, inner view; F—postabdominal claw; G— postabdominal seta; H–I—antenna I, external and anterior view. Scale bars 0.1 mm.

body rhomboid-rounded, thick, with a low and thick dorsal keel (Figure 1C). Moulting incomplete, reticulation on head shield and valves very fine and obscure. Head small, its ventral margin in posterior part with prominent basis for antennae I, labrum base surrounded with a low fold (Figure 1D). In ventral view, head shield triangular-ovoid, with prominent fornices, without a postero-lateral projection in region of process of mandibular articulation, the latter small (Figure 1E). Dorsal head pore located on a low prominence (Figure 1A, D-E, arrow). Compound eye of common size for genus, ocellus small, irregular in shape.



FIGURE 3. Ilyocryptus cuneatus brasiliensis subsp. nov., parthenogenetic female from Itaici, Municipio de Indaiatuba, São Paulo, Brazil: A–B—antenna II, anterior and external view; C—exopod; D—endopod; E—apical swimming seta of antenna II; F— proximal lateral seta; G— distal lateral seta; H— limb I; I–K— limb VI. Scale bars 0.1 mm.



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Labrum subquadrangular in lateral view, with a small medial projection in its basal portion and a prominent distal labral plate (Figure 1D). In ventral view, labrum body wide, with distinct paired projections in medial portion and levelled paired projections (supplied with two series of fine setules) in its distal portion (Figure 1F–G). Valves subovoid. Numerous setae along free margin (Figure 1H), five spaced, relatively short anteriormost setae, followed with a bunch of 4–5 closely located setae, which are just somewhat longer that following setae, the first seta in the bunch directed posteriorly (Figure 1I). Setae in middle of ventral margin with long setules, setae in postero-ventral region not longer that the former. Each seta at posterior margin along one side basally with series of spine-like setules, and distally with fine setules (Figure 1J–K). Abdomen dorsally with cross rows of setules (Figure 2A–C), a projection of size moderate for the genus on the first (basalmost) segment. Postabdomen relatively short, height maximal in the middle. Preanal margin somewhat longer than postanal one, with a row of regularly located, straight teeth, which are predominantly doubled. Few denticles near some of preanal teeth. Few denticles on lateral sides of postabdomen basally, or they completely absent. Anus small (Figure 2D), 5–6 spinules on its internal wall (Figure 2E). A row of relatively short paired spines starts on anal margin and continues up to distal boundary of preanal margin (Figure 2D). About seven large lateral setae longer than paired spines, the proximalmost lateral seta smaller, located on distalmost portion of preanal margin. On the distal part of postabdomen, the row of lateral setae fluently transits into a group of 4–5 middlesized setae, the latter, more distally, - into a group of rudimentary setae (Figure 2F). Postabdominal claw slightly bent (Figure 2F). There are three pectens of setules along concave dorsal margin. A single denticle on claw ventrally. Two spines of subequal size on the base of each claw dorsally. Long setules on claw base ventrally. Postabdominal seta longer than postabdomen, its basal segment approximately as long as distal one, the latter with short, sparse hairs (Figure 2G). Antenna I of medium length for Ilyocryptus, relatively thin. Bases of antennae I not compressed against each other (Figure 1F). Proximal segment with a well-expressed finger-like projection and low hillocks; distal segment without ridges and denticles (Figure 2H–I). Nine relatively short aesthetascs, two of them longer than the rest. Antenna II relatively short, coxal part with numerous relatively long setules, and two sensory setae of greatly differing size (Figure 3A–B). Distal sensory seta on basal segment long, slender, distal burrowing spine shorter than distal sensory seta, setulated distally. Antennal branches massive, on all segments, there are well-developed denticles around distal segment ends, and groups of similar denticles in middle part (Figure 3C–D). Swimming setae 0-0-0-3/1-1-3, spines 0-1-0-1/0-0-1. Apical swimming setae relatively short, bisegmented, differing in size, distal segments without hooks on tips, asymmetrically armed with minute setules (Figure 3E). Proximal lateral swimming setae shorter than distal one, both armed along one side with setules remarkably longer than those on apical setae (Figure 3F–G). Apical spine on exopod longer than spine on endopod. Spine on second segment of exopod about half of third segment length (Figure 3C). Limbs. Of armature typical of the genus. A large seta on outer distal lobe of limb I with basal segment armed unilaterally with sparse setules, and distal segment armed bilaterally with dense setules; a large, bisegmented, naked seta near ejector hooks, a gnathobase I as a naked lobe (Figure 3H). Limb VI as a trapeziumshaped plate with inner margin bearing continuous row of setules, separated by small incisions into six series, outer margin with series of setules (Figure 3I–K). Ephippial female, male. Unknown. Size. Parthenogenetic females from type locality 0.37–0.58 mm, holotype 0.55 mm. Distribution. At this moment, the taxon is known only from the type locality. Differential diagnosis. Ilyocryptus cuneatus brasiliensis subsp. nov. differs from the nominotypycal subspecies in: (1) smaller size, up to 0.58 mm; (2) presence of few additional small denticles near preanal teeth;



(3) reduced system of denticles on lateral faces of postabdomen; (4) fewer number of setules in bunch in middle portion of inner margin of limb VI. Comments. I. cuneatus cuneatus is the most common Palaearctic species, rarely occurring in the Nearctic zone (Kotov et al. 2002b; Kotov & Štifter 2006). Previously cuneatus-like forms were never described from any South of Central American countries. Here we demonstrate that two studied Neotropical populations belong to a separate taxon, although apparently closely related with Holarctic I. cuneatus cuneatus. Some differences revealed above are evidence of an aborigine status of this Brazilian population instead of an introduction from Holarctic due to a human activity. Some cuneatus-like populations are present in China, South Africa, Australia and new Zealand, but these forms must be revised in the future (Kotov & Štifter 2006).

Ilyocryptus silvaeducensis paraensis subsp. nov. Figure 4 Undescribed subspesies of I. silvaeducensis in Kotov and Štifter, 2005a: p. 8, Figs 35–76; Kotov and Štifter, 2006: p. 125, Fig. 62N.

Etymology. This subspecies is named after the state of Brazil where it was found, Pará. Type locality. Lake Batata, Porto Trombetas (01°30'0.00''S, 56°19'30.34''W), Município de Oriximiná, Pará, Brazil. The type series was collected in xii.1988 by R.L. Bozelli. Type material. Holotype. A juvenile parthenogenetic female, MNRJ 21505. The label of holotype is: "Ilyocryptus silvaeducensis paraensis subsp. nov., 1 parth. & from Lake Batata, Pará, Brazil, HOLOTYPE". Paratypes. 2 juvenile females, MGU Ml 76. Short description. Adult parthenogenetic female. See Kotov and Štifter (2005a). Juvenile female. General: In lateral view body triangular-ovoid, relative height of juveniles small for the genus (body height/body length = 0.66–0.70 in juveniles), maximum height in posterior half (Figure 4A). Dorsal margin straight, postero-dorsal angle expressed. In anterior view, body compressed laterally, with distinct, sharp dorsal keel (Figure 4B), but we do not know, is this characteristic of adults also? Molting incomplete, but first instar female has no previous exuvia on valves and head shield (Figure 4A). Reticulation on head shield and valves very fine. Head small, its ventral margin in posterior part with a low basis for antennae I (Figure 4C). Valves subovoid. Numerous setae along free margin, five anteriormost setae protruding sparsely and very long (this is a juvenile character, in adults these setae significantly shorter), followed with a bunch of 5–7 closely located setae, which are only somewhat longer that following setae, the first seta in bunch directed posteriorly. Setae in middle of ventral margin with long setules, setae in postero-ventral region not longer that the former. Each seta at posterior margin basally with series of spine-like setules along one its side (Figures 4E–D). Postabdomen relatively short, height maximal in middle. Preanal margin approximately as long as postanal one (this is a juvenile character, in adults preanal margin shorter), with a row of 5–6 regularly located, straight teeth, which are predominantly or exclusively doubled, 1–2 distalmost teeth small (Figure 4F–G). Rare denticles near preanal teeth (a juvenile character, in adults these are absent). Few denticles on lateral faces of postabdomen basally. Anus small, 7–9 spinules on its internal wall (Figure 4H). A row of relatively short paired spines starts on anal margin and continues up to distal boundary of preanal margin. Large lateral setae longer than paired spines, the proximalmost lateral seta located on distal portion of preanal margin, several lateral setae present at anal margin. On the distal part of postabdomen, the row of lateral setae fluently transits into the group of 4–7 middle-sized setae, the latter, more distally, – into group of rudimentary setae.



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FIGURE 4. Ilyocryptus silvaeducensis paraensis subsp. nov., juvenile female from Lake Batata, Pará, Brazil: A–B—lateral and anterior view; C—head, lateral view; D–E—setae at ventral and dorsal portion of posterior margin; F–G—postabdomen; H—anus, inner view; I—antenna II; J–K—apical and lateral swimming seta. Scale bars 0.1 mm.

Postabdominal claw slightly bent. Few denticles on claw ventrally. Distalmost spine on base of postabdominal claw slightly longer than proximal one. Long setules on claw base ventrally. Postabdominal seta longer than postabdomen, its distal segment with long, sparse hairs (Figure 4A).



Antenna I of medium length for Ilyocryptus, bases of antennae I not compressed against each other. Proximal segment with a well-expressed finger-like projection and low hillocks; distal segment without ridges and denticles, distal end with concentric row of small hillocks. Nine relatively short aesthetascs, two of them longer than the rest. Antenna II relatively short, coxal part with two sensory setae of greatly differing size (Figure 4I). Distal burrowing spine short, reaching only the distal end of basal segment. Antennal branches massive, on all segments, there are well-developed denticles around distal segment ends, and groups of similar denticles in middle part. Swimming setae 0-0-0-3/1-1-3, spines 0-1-0-1/0-0-1. Apical swimming setae asymmetrically armed with minute setules (Figure 4J). Lateral swimming setae armed along one side with setules on type of apical setae, and along another side with remarkably longer setules (Figure 4K). Apical spine on exopod longer than spine on endopod. Spine on second segment of exopod longer than half of third segment. Limbs. Typical of the genus. Ephippial female, male. Unknown. Size. Juvenile females from type locality 0.29–0.31 mm, holotype 0.31 mm. Maximum size is greatly underestimated due to absence of adults in our sample. Distribution. At this moment, the taxon is known only from two localities in northern portion of Brazil: (1) Lake Batata, Porto Trombetas, Pará and (2) a small, shallow pool near Lagoa da Colher, the eastern border of the Lençóis Maranhenses dune field, Maranhão. Probably, this is a rare species in Brazil. Differential diagnosis. Ilyocryptus silvaeducensis paraensis subsp. nov. differs from two other known subspecies in: (1) presence of several lateral setae at anal margin; (2) relatively short distal burrowing spine, not protruding beyond the distal end of basal segment of antenna II; (3) specially thick segments of branches of antenna II (Kotov & Štifter 2005a, 2006). Comments. Only three juvenile females were found, and one of them was designated as the holotype. I. silvaeducensis silvaeducensis occurs only in North Europe, the Nearctic zone is populated by a separate subspecies I. silvaeducensis chengalathi. Here we establish a new Neotropical subspecies. So, subspecies of I. silvaeducensis conform to main biogeographical zones. Kotov and Štifter (2005a) found that Brazilian subspecies (un-named at that time) is more distant from European and North American subspecies, which are closest congeners. African (Kotov 2000) and Australian (Sars 1896) silvaeducensis-like populations must be specially examined in the future.

Ilyocryptus paranaensis paranaensis Paggi, 1989 Figure 5 Daday 1905: 191 (Iliocryptus sordidus); Paggi 1989: 240–245, Fig. 1–39; Kotov et al. 2001: 1069–1070; Kotov and Stifter 2005b: 110–112, Figs 1–11; Kotov and Štifter 2006: 110, Figs 53–54.

Type locality. Unnamed, weedy oxbow at Island Los Sapos, in the lower course of the River Salado near Santo Tomé, Paraná River basin, Province of Santa Fe, Argentina (Paggi 1989). Material examined here. An adult parthenogenetic female 0.69 mm from Rio Atibaia between Itatiba and Campinas (appr. 22º57'S, 46º50'W), São Paulo, coll. on 29.iii.2000 by M.L. Kuhlmann; an adult parthenogenetic female from Lagoa do Pesqueiro, Itaici, (23°06'48.2" S, 47°11'03.0"W), Municipio de Indaiatuba, São Paulo, coll. on 14.xi.2002 by L. M. A. Elmoor-Loureiro; a parthenogenetic female from Parque Ecológico do Tietê, São Paulo city (appr. 23°29'S, 46°31'W), São Paulo, coll. on 05.iii.2001 by F. Foratto. Diagnosis of nomenotypical subspecies. Cervical incision shallow (Figure 5A–B). Spinules at valve margin on inner surface (Figure 5C). Denticles on base of postabdomen well developed (Figure 5D). Proximal segment of antenna I smooth, distal segment with obscure transverse ridges. Spine of second segment of antennal exopod look unsegmented, with rounded tip, without setules (Figure 5A). REVISION OF ILYOCRYPTUS OF BRAZIL


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Description. See Kotov and Štifter (2005b). Comments. This species is recorded for the first time for Brazil. Earlier I. paranaensis paranaensis was found in Argentina, Paraguay and Venezuela, while I. paranaensis inarmatus inhabits the Usumacinta basin, Mexico (Paggi 1989; Kotov et al. 2001; Kotov & Štifter 2005b, 2006).

FIGURE 5. Ilyocryptus paranaensis paranaensis, adult parthenogenetic female from Atibaia River, between Itatiba and Campinas, São Paulo, Brazil: A–B—lateral and dorsal view; C—posterior margin of valve, inner view; D—preanal portion of postabdomen. Scale bars 0.1 mm.

Ilyocryptus sarsi Stingelin, 1913 Sars 1901: 42, Pl. 7: Figs. 11–13 (Ilyocryptus sordidus); Stingelin 1913: 618 (Iliocryptus sordidus var. sarsi); Smirnov 1976: 49, fig. 14 (Ilyocryptus sordidus sarsi); Kotov et al. 2002a: 208–219, figs 1–60; Kotov and Štifter 2006: 118–123, Figs 58–60.

Type locality. Ipiranga, State of Sno Paulo, Brazil (according to neotype). Neotype. An adult female, GOS F18459a. Material examined here. Several females from neotype locality, GOS F18459b; four parthenogenetic females from Billings Reservoir, São Paulo city, São Paulo, coll. on 14.iii.2001 by M.J.C. Botelho; two parthenogenetic females from Parque Nacional da Serra do Cipó (19° 20' 55" S; 43° 36' 32" W), Minas Gerais, coll. on 30.v.1997 by A.O. Bustamante; 27 parthenogenetic females from Mucugezinho River, Lençóis, Bahia, coll. on 30.v.1997 by V. M. G. Barros; a parthenogenetic female from Lagoa Grande, Balsas, Maranhão, coll on 15.vi.2001 by M.S. Ibañez. Diagnosis, description. See Kotov et al. (2002a). Comments. This is a relatively common species.

Ilyocryptus spinifer Herrick, 1882 Herrick 1882a: 246, Pl. 8: figs 2–6; Herrick 1882b: 39–41, Pl. 9: figs 1–3; Ihering 1895: 168, figs 1–2 (Acanthocercus immundus Fr. Mueller); Sars 1901: 40–42, Pl. 7: figs 1–10 (longiremis); Bergamin 1939, fig. 4. (Iliocryptus tetraspi-



natus); Brandorff et al. 1982: 94, Figs 67–69; Kotov and Dumont 2000: 88–105, figs 1–149; Hollwedel et al. 2003: 78; Elmoor-Loureiro 2004: 54, figs 1–10; Kotov and Štifter 2006: 139–144, Figs 1, 3, 6–7, 9–11, 69–70. (Only first Herrick’s descriptions and papers concerning Brazil are listed).

Type locality. A small bog located in an inlet stream of Lake Alice, Hubbard County, Minnesota, U.S.A. (according to neotype selected by Kotov and Williams, 2000). Material examined here. Roraima: near BR 174 road, between Boa Vista and Mucajaí, coll. on 31.vii.2007 by L.F.R. Guterres. Pará: Lake Batata, Porto Trombetas (01°30'0.00''S, 56°19'30.34"W), coll. on xii.1988 by R.L. Bozelli; pool at Museu Paraense Emilio Goeldi (appr. 01°27'S, 48°28'W), Belém, coll. on 31.x.1999; pond at Serra dos Carajás (06°24'30"S, 50°21'04"W) coll. R.L. Bozelli. Maranhão: Turiaçu River, Santa Helena (appr. 02°13'41"S, 45°18'05"W), coll. on 05.v.1999 by R. Barbieri & J.P. Costa Neto; Lago Viana, dowstream River Cajari, (appr. 03°13'S, 45°00'W), coll. on 06.v.1999 by R. Barbieri & J.P. Costa Neto. Ceará: well marginal to Lagoa Precabura, Fortaleza (appr. 03°48'S, 38°26'W), coll. on 12.vii.2000; pond marginal to Lagoa do Banana, Caucaia (appr. 03°37'S, 38°45'W), coll. on 14.vii.2000. Pernanmbuco: pond at BR232 road (appr. 08°12'S, 35°34'W), Muncípio de Gravatá, coll. on 27.ii.1995 by P.S. Young & C.S. Serejo; Açude de Dois Irmãos (08°00'51"S, 34°56'47"W), Recife, coll. on 17.i.2000. Bahia: Pituaçu reservoir (appr. 12°58'1"S, 38°24'44"W), Salvador, coll. on 04.viii.1981 by J.W. Reid; Lagoa do Abaeté (12°56'44"S, 38°21'27"W), Salvador, coll. on 28.vii.1985; pond at BR020 road (appr. 12°47'18"S; 45°57'06"W), Roda Velha, Minicípio de São Desidéio, coll on 30.vii.1985; temporary pool at Km 860 of the BR101 road, Município de Teixeira de Freitas (17°32'S, 39°44'W) coll. on 15.ii.1997 by P.S. Young & C.S. Serejo; Capivari River, Muncípio de Cruz das Almas (appr. 12°39'S, 39°07'W), coll. on 09.x.2000 by E.M. Silva; pond at Praia do Forte (12°34'41"S, 38°00'47"W), Muncípio de Mata de São João, coll. on 12.vii.2002; pool near de airport, Morro de São Paulo (13°25'49”S, 38°54'48"W), Muncípio de Cairu, coll. on 29.i.2004; Lagoa do Formoso, Trijunção Farm (14°49'02"S, 45°58' 35"W), Muncípio de Cocos, coll. on 26.iv.2003. Goiás: pool near Pirenópolis (appr. 15°51'8"S, 48°57'58"W), coll. on 15.ii.1988; pond at Guizzetti Farm, BR153 road (18°20'34"S, 49°13'14"W), Muncípio de Itumbiara, coll. on 07.v.2000; farm reservoir (15°45'12"S, 48°40'07"W), Município de Cocalzinho de Goiás, coll. on 30.ix.2000. Distrito Federal: Lagoa Bonita (15°35'22"S, 47°41'50"W), Planaltina, coll. on 07.x.1980; Lagoa Joaquim Medeiros (15°38'16"S, 47°41'30"W), Planaltina, coll. on 08.viii.2006; pond at Chacara Ita (15°40'33"S; 47°41'37"W), Planaltina, coll. on 08.viii.2006; Lagoa Taquara (15°38'12"S, 47°31'22"W), Planaltina, coll. on 15.viii.2006; Sobradinho River (15°39'20"S, 47°48'55"W), Sobradinho, coll. on 18.viii.2006; Corrego Tabocas (15°52'29.5''S, 47°43'34.1''W), coll. on 06.ix.2006 by F.D. Sousa; Lagoa do Henrique (15°41'16"S, 47°56'22"W), Parque Nacional de Brasília, coll. on 27.v.2002; Lagoa do Cedro (15°53'50.2"S, 47°56'37.7"W), Brasília, coll. 14.i.2006. Mato Grosso: swamp near Poconé (appr. 16°25'31"S, 56°40'8"W), coll. on 06.ii.1986; swamp near Santo Antônio do Leverger (appr. 15°50'26"S, 56°04'33"W), coll. on 26.vii.1988; Guaporé River (appr. 14°00'42"S, 59°56'46"W), Vila Bela da Santíssima Trindade, coll on xii.2006, leg. E. Porto. Mato Grosso do Sul: baía at Fazenda Nhumirim (18°59'34.9"S, 56°35'59.9"W), Município de Corumbá, coll. on 04.x.2001 by V.M.G. Barros; fish pool at Centro Laura Vicuña, Campo Grande (appr. 20°31'S, 54°39'W), coll. on 06.viii.2000. Minas Gerais: Lagoa Cajueiro and Lagoa Juazeiro (appr. 15°05'S, 44°03'W) Município de Itacarambi, coll. on iii.1996 by M.B. G.Sousa; Preto River (appr. 18°03'S, 43°22'W), Município de São Gonçalo do Rio Preto, coll. on iii.1996 by M.B.G.Sousa; pond at Caparó Parque Hotel (appr. 20°25'S, 41°51'W), Município de Alto Caparó, coll. on 27.i.1991; Tanque São Luiz and pond near the hostel at Parque Natural do Caraça (appr. 20°05'S, 43°29'W), Município de Santa Bárbara, coll. on 22.vii.1992; Parque Nacional Grande Sertão Veredas (15°16'12"S, 45°56'44"W), Município de Chapada Gaúcha, coll. on 19.iii.2001 by G. Miranda. Espírito Santo: Lagoa Juparanã (appr. 19°18'S, 40°05'W), Município de Linhares, coll. on 25.i.1991. Rio de Janeiro: Lagoa Cabiúnas (22°18’S, 41°42’W), Município de Macaé, coll. on iv.2004 by P. Lopes; Município de Saquarema (appr. 22°56'S, 42°29'W), coll. 18.iv.1998 by J.W. Reid & P.S. Young. São Paulo: reservoir at Santana de Baixo Farm (appr. 22°17'S, 47°41'W), Município de Corumbataí, coll. on 24.viii.1996; Lake REVISION OF ILYOCRYPTUS OF BRAZIL


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Itapeva (22°46'25S,45°33'24”W), Campos do Jordão, coll. on 15.xi.2002; Paranapanema River and its marginal lagoons (appr. 23°29'S, 48°37'W), Município de Angatuba, coll. on 21.x.2000; Tietê River (appr. 23°34’S, 46°02'W), Município de Biritiba Mirim, coll. on 13.ii.1993 by M.L. Kuhlmann; Ponte Nova Reservoir (appr. 23°34'S, 45°58'W), Município de Biritiba Mirim, coll. on 02.xii.1997 by M.J.C. Botelho; Lagoa do Pesqueiro, Itaici, (23°06'48.2" S, 47°11'03''W), Municipio de Indaiatuba, São Paulo, coll. on 14.xi.2002. Paraná: reservoir at Fragosos (appr. 26°05'S, 49°25'W), Minicípio de Pien, coll. on 20.xi.1999 by P.S. Young & C.S. Serejo. Santa Catarina: fish pond at Mapiju Farm, Município de Santo Amaro da Imperatriz (appr. 27°41'S, 48°46'W), coll. on 27.ix.2001 by G. Pereira Jr. Rio Grande do Sul: Lago do Parque Knorr (29°22'45"S, 50°52'02"W), Gramado, coll. on 31.xii.2001; Lago Negro (29°23'44"S, 50°52'33"W), Gramado, coll. on 03.i.2002; Lago Guaíba (30°08'33"S, 51°13'36"W), Porto Alegre, coll. on 27.xii.2001; Lago Jacaré (32°36'03"S, 52°34'03"W) and Canal de Sarita (32°37'29"S, 52°35'48''W), Estação Ecológica do Taim, Município de Rio Grande, coll. on 29.xii.2001. Diagnosis, description. See Kotov and Dumont (2000), where many Brazilian specimens were illustrated. Comments. This is the most common Brazilian species.

Discussion The Neotropical fauna of the Ilyocryptidae included nine known species (Forró et al. 2008) before our finding of two new taxa. We found only five species of Ilyocryptus having analysed many (hundreds) samples from Brazil (Figure 6). Among them three taxa are exclusively South American (I. cuneatus brasiliensis subsp. nov., I. silvaeducensis paraensis subsp. nov., and I. paranaensis paranaensis). I. spinifer has a very wide distributional area (Kotov & Dumont 2000). “I. sarsi” seems to be a group of close topical congeners (Kotov & Štifter 2006); quite possible that I. sarsi s. str. is an exclusively Neotropical taxon. So, Neotropical endemics dominate among Brazilian ilyocryptids. We believe that also there is a chance to find I. elegans Paggi, 1992 and I. plumosus Kotov & Štifter, 2004 in tropical lowlands. The latter taxa are also endemics of South America. Most part of records of Brazilian ilyocryptids concerns I. spinifer, which is among the most common cladocerans in the Neotropics (Kotov & Štifter 2006). Other ilyocryptid taxa are significantly rarely occur, although it is necessary to note that their absence in samples may be partly explained by inadequate methods of collections. Most investigators collect only plankton and the fauna of vegetation zone, where ilyocryptids are absent or rare (I. spinifer and few other species are exceptions from this rule, see Kotov and Štifter 2006). I. sordidus (Lévin, 1848), reported previously by many authors from different Neotropical localities, was not found in our material from Brazil. We believe that previous records were misidentifications, dealt predominantly with two aforementioned members of I. silvaeducensis-group, or with I. sarsi. Unfortunately, most part of previous records of “I. sordidus” from Brazil, as well as other South American countries, could not be attributed to any real species, because these papers did not contain any descriptions or illustrations. This is, in reality, a “dead” faunistic information, useless for any kinds of subsequent analysis. The main our discovery is finding of two new subspecies of species earlier described from Palaearctic, namely I. cuneatus and I. silvaeducensis. Establishing separate subspecies, not species, for them we specially underline closest relations of these Palaearctic and Neotropical forms. But both I. cuneatus and I. silvaeducensis have unrevised populations from other continents, so, our finding cannot be interpreted as an evidence of especial Europe-South American biogeographical relationships. A final conclusion can be made only after a global revision of cuneatus and silvaeducensis-like populations worldwide. In case of I. cuneatus, which is the most common Palaearctic species, isolated populations of its congeners in southern hemisphere could be regarded as relicts sensu Korovchinsky (2006). Their existence suggests an earlier differentiation within the



FIGURE 6. Distribution of four Ilyocryptus species (A) and Ilyocryptus spinifer (B) in Brazil, from original (solid symbols) and literature data (open symbols).



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cuneatus-group, confirming ideas of earlier differentiation of the cladoceran taxa (Fryer 1995; Korovchinsky 2006; Kotov 2007). The overall cladoceran species richness is probably up to 4 times higher than currently known (Adamowicz & Purvis 2005; Forró et al. 2008). Kotov and Štifter (2006: p. 5) noted that “we can confidently say that the species diversity within the genus Ilyocryptus remains underestimated and that a series of new species will be discovered and described, as revisions of some species groups will progress in the future”. This issue is now confirmed for Brazil.

Acknowledgements We are very grateful to Prof. N.N. Smirnov for valuable consultations. We would like to thank the following persons for providing specimens: Eduardo M. da Silva, Eucilene Porto, Guilherme Miranda, Gilberto Pereira Jr, Janet W. Reid, Luis F.R. Guterres, Márcia J. C. Botelho, Maria Beatriz G. Souza, M. Socorro Ibañez, Reinaldo L. Bozelli, Ricardo Barbieri, Valéria M.G. Barros, and Paulo S. Young (in memorian). We also thank Rodrigo G. Gonçalves for his assistance in the maps preparation. This work is supported by Russian Foundation for Basic Research (grant 06-04-48624), "Biodiversity" Program of the Presidium of Russian Academy of Sciences (grant 1.1.6) and Russian Science Support Foundation (for AAK) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (grant 555288/2006-4 for LMAEL).

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