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Abstract
Emended diagnoses of all stages of the genus IchthyocladiusFittkau are given. The genus appears to be a basal memberof the Corynoneura group showing affinities also to the Euki-efferiella group. A number of features apparently are theresult of the phoretic life on catfishes. Ichthyocladius kro-nichticola sp. n. and I. lilianae sp. n. are described andfigured in all stages. Additional pupae from the Amazonas inBrazil and from the Parque Nacional Iguazú in Argentina arealso described and figured. Keys to pupae and larvae of thegenus are given.
Keywords: Chironomidae, Orthocladiinae, Corynoneura –group, Ichthyocladius, new species, Brazil, Argentina.
Introduction
Fittkau (1974) erected the genus Ichthyocladius for twospecies from Peru and Ecuador. The larvae of the genus livephoretically on catfishes of the families Astroblepidae andLoricariidae, attaching themselves to the skin and scales ofthe fish. At the end of the fourth larval stage the larvae spina holdfast that is attached to spines or fin rays and finallyformed into a pupal case. The only named species, I. neotrop-icus Fittkau, was described based on pharate pupae and asso-ciated larvae. Several details, especially concerning thewings and the apodemes of the male hypopygium, thus havenot been described previously.
Fittkau (1974) mentioned several larvae of the genus from Trinidad, Venezuela, Guyana, Bolivia and Brazil not identified to species. In the present paper all stages of twonew Brazilian species and the pupal exuviae of two addi-tional species from Brazil and Argentina are described andfigured.
Material and methods
The general terminology follows Sæther (1980). The malehypopygium is figured with tergite IX removed, dorsal aspect to the left, and ventral aspect to the right. Dorsal view of tergite IX, gonocoxite and gonostylus are figuredseparately.
All material is mounted on microscope slides and prepared in Canada Balsam or Euparal. The holotypes and paratypes of the new species are deposited at the Museu de Zoologia da Universidade de São Paulo, Brazil(MZUSP). Paratypes are also deposited at the ZoologischeStaatssammlung, Munich, Germany (ZSM), and at theMuseum of Zoology, University of Bergen, Norway(ZMBN).
Ichthyocladius Fittkau
Ichthyocladius Fittkau 1974: 91.
New species of Ichthyocladius Fittkau, a Member of the
Corynoneura-Group (Diptera: Chironomidae: Orthocladiinae),
with a Review of the Genus
Humberto F. Mendes1, Trond Andersen2 and Ole A. Sæther2
1Department of Biology-FFCLRP, University of São Paulo, Ribeirão Preto, Brazil2Museum of Zoology, University of Bergen, Bergen, Norway
Received: 19 January 2003Accepted: 21 November 2003
Correspondence: T. Andersen, Museum of Zoology, University of Bergen, Muséplass 3, N-5007 Bergen, Norway. Fax: +47 55589677; E-mail: [email protected]
Studies on Neotropical Fauna and Environment2004, Vol. 39, No. 1, pp. 15–35
DOI: 10.1080/01650520412331270936 © 2004 Taylor & Francis Ltd.
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16 H.F. Mendes et al.
Type species
Ichthyocladius neotropicus Fittkau, 1974, by original desig-nation and monotypy.
Other included species
Ichthyocladius kronichticola sp. n.; Ichthyocladius lilianaesp. n.; Ichthyocladius sp. Ecuador; Ichthyocladius sp. RioMarauiá, Brazil; Ichthyocladius sp. Argentina.
Diagnosis
Imago. Small species, wing length about 1.3mm.Eye bare, rounded, no dorsomedian elongation. Antenna
with 12 (or perhaps sometimes fewer, see below) flagellom-eres in male, six in female; male antenna sparsely plumed;groove beginning on flagellomere 3; sensilla chaetica minute,present on flagellomeres 2, 3 and 12; apex without straightapical seta; AR lower than 0.4. Only one palpomere, no sen-silla clavata. Temporals consisting of a few minute inner andouter verticals (or sometimes absent?). Tentorium with broadbut short base, stipes apparently normal. Cibarial pumpbroad, but short, with concave anterior margin and welldeveloped, triangular cornua. Clypeus narrow with shortlateral setae.
Antepronotal lobes broad, collar-like, not medially nar-rowed, without setae. Scutal tubercle present and prominent.Acrostichals absent, dorsocentrals uni-triserial, several pre-alars, supraalar present. Scutellum with setae transverselyuniserial.
Wing broad, with well developed, but not projecting anallobe; membrane with distinct punctation; costa slightlyextended; veins R1 and R2+3 short, thick and fused with costain thick clavus, ending slightly beyond midpoint of wing;R4+5 weak; VR moderately high; Cu1 slightly sinuous; postcu-bitus ending distal to cubital fork, anal vein ending below orslightly distal to cubital fork. Brachiolum with one seta, otherveins bare. Squama bare. Sensilla campaniformia about eightbasally and eight apically on brachiolum, apparently nonebelow setae on brachiolum; one present basally on subcosta,and one on RM.
Front leg ratio high, about 0.8. Front tibia with straight,weak spine-like tibial spur; spurs of mid tibia either withthorn-like lateral denticles or entirely consisting of thorn-likespines; hind tibia with well developed spurs covered withthorn-like lateral denticles; hind tibial comb absent orreduced to a few setae. Short tarsal pseudospurs present onmetatarsi of mid and hind legs and ta2 of mid leg. Pulvilliabsent. No sensilla chaetica observed.
Tergites II–VII in male, II–VIII in female each with groupof median setae sometimes on a low protuberance, tergitesexcept for male tergite IX otherwise bare; sternites except forfemale sternite VIII bare.
Tergite IX of male large, covering much of gonocoxites,with notched or bilobed posterior margin, with several apical
and some apical ventrolateral setae; laterosternite IX bare.Sternapodeme strong, triangular. Phallapodeme and aedeagallobe normally developed. Virga absent. Gonocoxite welldeveloped, superior volsella apparently absent, inferiorvolsella low to moderately well developed. Gonostylusbroadest near apex, without crista dorsalis and megaseta.
Female genitalia with well developed gonocoxite withseveral setae. Tergite IX very large, covering most of geni-talia, medially divided, with several apical setae. Segment Xapparently normal. Postgenital plate weak or absent. Cercustriangular, small. Gonapophysis VIII undivided. Shape ofapodeme lobe and coxosternapodeme unknown. Seminalcapsules much larger than cerci, ovoid; with orally placedneck. Spermathecal ducts nearly straight, apparently withcommon opening. Labia not known.
Pupa. Small pupae, about 2–3mm long. Exuviae nearlytransparent to strongly marked with brown.
Cephalic tubercles and frontal warts absent. A low protu-berance densely covered with tubercles present basally onpedicel. Frontal setae absent, frontal apotome wrinkled. Tho-racic horn absent. Wing sheath smooth, without pearls ornose, but sometimes with notch. Thorax with median tuber-cle. Three precorneals, four antepronotals, at least one pos-torbital, and four dorsocentrals present, all minute.
Segment I and sternite IX without shagreen; tergites andsternites II–VIII with shagreen, on tergites II–VII medianshagreen spinules stronger and sometimes placed on protu-berance. Conjunctives bare. Tergites II–V and sternitesIV–VII each with 34–50 caudal hooklets; a few lateral ormedian hooklets sometimes present also on tergite I and/orsternite III. Pedes spurii A and B absent.
Abdominal setae minute; segments I–V each with 1Lseta, VI–VII with 1–2L setae, VIII with four L setae; tergitesI–VI with four, VII with three and VIII with one pair of Dsetae; sternite I with three, II with four, III–VII with five, andVIII with two pairs of V setae. Segment IX with three short,scimitar-like or hair-like curved macrosetae situated ven-trally or at apex. O setae not observed. Genital sac of maleslightly to much longer than anal segment, broadly rounded,without shagreen or apically wrinkled and with a few minuteapical spinules.
Larva. Small larvae, up to 7mm long.Antenna with four segments, basal segment about as long
as or slightly longer than flagellum, second antennal segmentslightly longer than or subequal to third. Basal antennalsegment 1.5–2 times as long as basally wide, with ring organsituated in apical third. Lauterborn organs not observed, stylewell developed. Blade slightly shorter than flagellum, acces-sory blade about half as long.
S I pectinate to plumose. Other S setae simple. Labrallamella large, bluntly triangular with about 50 teeth. Chaetaereduced to two pegs. Pecten epipharyngis consisting of threepointed teeth. About six chaetulae laterales with the median
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New Species of Ichthyocladius 17
apparently serrated, chaetulae basales weak. Premandiblewith three distinct dark inner teeth and 3–5 broadly lamel-late, pale and fused outer teeth (not visible in all views),without or with vestigial brush.
Mandible with long, bent apical tooth longer than com-bined width of 3–4 inner teeth. Seta subdentalis well devel-oped. Seta interna absent.
Mentum with 16–20 equal-sized small median teethnearly in a straight line and five pairs of slightly larger lateralteeth. Ventromental plates long and narrow. Setae submentisituated above base of outer lateral tooth of mentum.
Maxilla relatively large, pecten galearis apparently absent,maxillary palp well developed. Anterior lacinial chaetaapparently with apical teeth.
Anterior and posterior parapods well developed; claws ofanterior parapods pale to brown, longer claws smooth or witha few very small inner teeth; posterior parapods with 12–15black claws without teeth. Procercus absent or very low, 3–4short anal setae. Supraanal seta minute. Anal tubules veryshort. Body setae very short.
Diagnostic characters
The male imagines are separable from all other chironomidsby the presence of a clavus combined with a bare scutaltubercle and the lack of a gonostylar megaseta. The pupaediffer from all chironomids by the possession of hooklets ontergites I–IV and sternites IV–VII combined with lack of athoracic horn and curved (associated pupae) or hair-like (un-associated pupae) macrosetae mostly on the ventral sideof the anal lobe. The larva differs from other chironomids by having about 16–20 dark, equal-sized median and fivepairs of slightly larger lateral teeth in the mentum and nomandibular seta interna.
Systematics
The presence of a clavus places the genus near CorynoneuraWinnertz and related genera, while the caudal hooklets of thepupa combined with the absence of a fringe on the anal lobeindicate relationship with the Eukiefferiella (Cardiocladius)group. However, the two groups may be sister groups.
The presence of large caudal hooklets on the tergal con-junctives or posterior on the tergites is conspicuous both inIchthyocladius, Tempisquitoneura Epler and a new genusnear Thienemanniella Kieffer and every bit as conspicuousas those in the Eukiefferiella group. Sæther (1977) placed theCorynoneura group as sister group to a reduced Orthocladi-ini sensu Brundin and a Metriocnemini sensu Brundin com-bined, i.e., with some plesiomorphic groups removed, withHeterotanytarsus Spärck as sister group to all three groupscombined and the Eukiefferiella group as sister to all theabove. That is, the Corynoneura group was just two nodesremoved from the Eukiefferiella group. The new genus nearThienemanniella, Tempisquitoneura and Ichthyocladius,however, strengthen the indications of relationship.
Characters in common for all the genera of the two groupsinclude absence of pedes spurii A and B of the pupa; sexdimorphism of sternite VIII of the pupa; gonapophysis VIIIof the female genitalia undivided; and ventromental plates ofthe larvae narrow, inconspicuous, widest posteriorly andwithout setae underneath.
Ichthyocladius, in addition to the possession of a clavus,shares the reduced temporals, the large tergite IX coveringmost of the male hypopygium, absence of anal point andvirga, seminal capsule with orally placed neck, and nearly straight spermathecal duct with Corynoneuraand Thienemanniella Kieffer. An inverted V-shaped ster-napodeme without oral projection is present also inCorynoneura. However, Ichthyocladius also has several fea-tures not found in Corynoneura and Thienemanniella. Someof these, such as the reduced larval and pupal features andthe absence of a megaseta on the male gonostylus, likely area result of the phoretic life style, while others appear to beplesiomorphies. In the last group are non-cordiform tarsi,presence of tarsal pseudospurs, and the well developed anallobe and distinct punctations of the wings. That the clavusends in the distal half of the wing instead of in the basal halfalso appears to be a plesiomorphy as compared to othergenera of the Corynoneura group (Sæther & Kristoffersen,1996).
Other characteristic features include the scutal tuberclewithout any microtrichial tuft or acrostichals which other-wise amongst the Orthocladiinae occurs only in RheosmittiaBrundin. The one-segmented palp otherwise is found only insome species of Symbiocladius Kieffer (Gonser & Spies,1997). At first the pupa was thought to lack setae, spines orthorns from the pupal anal segment as otherwise found onlyin very few genera or species including Rheosmittia andEukiefferiella ancyla Svensson, a commensal in Ancylus flu-viatilis Müller (Svensson, 1986). However, there are threemacrosetae mostly placed ventrally on the anal lobe betweenthe lobe and the genital sac and a minute additional apico-lateral seta. These setae may easily have been overlooked alsoin other genera. Fittkau (1974: fig. 11) is incorrect in describ-ing the S setae, S I really is S II and S II is S III, while S Iis figured as part of the pecten epipharyngis. Thus S I is infact pectinate or plumose, a feature found in Tvetenia Kiefferof the Eukiefferiella group, but not in genera nearCorynoneura. The labral lamella is large, bluntly triangularand finely pectinate or denticulate, but often is perpendicu-lar to the labral surface and thus obscured. A large labrallamella is not found in any genus near Corynoneura or Euki-efferiella. S I appear to be attached underneath the labrallamella, i.e., about as in Metriocnemus v. d. Wulp (Cranstonet al., 1983: fig. 9.43).
The larval mentum is of the same type as the mentum ofsome other phoretic larvae such as Baeoctenus Sæther, Drat-nalia Sæther et Halvorsen and Epoicocladius Zavrel. Sæther(1986) discuss the menta of these genera as a prime exampleof parallel selection and suggests that the menta act as combsscraping off particles from hairs and appendages of their
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hosts. Of the above genera, Dratnalia belongs in the Eukief-feriella group. Amongst genera related to Corynoneura, Tem-pisquitoneura is known to be symphoretic on Corydalus(Megaloptera: Corydalidae) (Epler & De la Rosa, 1995).However, in that genus the immatures are normal.
Key to known pupae of Ichthyocladius Fittkau
1. Tergites II–VII with strong median shagreen spinulesplaced on protuberance. Brazil (Figs. 11–15) . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. kronichticola sp. n.– Tergites II–VII without protuberance or protuberance atmost indicated on tergite VI and VII . . . . . . . . . . . . . . . . 22. Tergite I without posterior spines or hooklets, hooklets on tergite II distinctly weaker than those on tergites III–IV, exuviae pale. Peru (Fittkau, 1974: figs. 4 & 5) . . . . . . . . . . . . . . . . . . . . . . . . . . I. neotropicus Fittkau– Tergite I with weak posterior hooklets, hooklets on tergiteII as strong as those on tergites II–IV . . . . . . . . . . . . . . . 33. Exuviae pale; anal macrosetae strongly curved, scimitar-like, broad. Brazil (Figs. 23–25) . . . . . . . . I. lilianae sp. n.– Exuviae amber with dark brown margins; anal macrosetaehair-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44. Abdominal apex rounded, gradually narrowed betweensegment VIII and IX, female genital sac ending far short ofapex of anal segment. Brazil (Figs. 32, 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sp. Rio Marauiá– Abdomen abruptly narrowed between segment VIII andIX, male genital sac much longer than anal segment and con-spicuously bicolored, female genital sac reaching apex ofanal segment. Argentina (Figs. 34, 35) . . . . I. sp. Argentina
Key to known larvae of Ichthyocladius Fittkau
1. Mandible with three inner teeth. Ecuador (Fittkau, 1974:fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sp. Ecuador– Mandible with four inner teeth . . . . . . . . . . . . . . . . . . 22. Procercus absent, three anal setae. (Fittkau, 1974: fig. 13) . . . . . . . . . . . . . . . . . . . . . . I. neotropicus Fittkau– Procercus present, four anal setae . . . . . . . . . . . . . . . . 33. Mentum with 16–18 median teeth and five pairs of lateral teeth (Figs. 16–20) . . . . . . . . . . . . . I. kronichticola sp. n.– Mentum with 20 median teeth and five pairs of lateral teeth(Figs. 26–31) . . . . . . . . . . . . . . . . . . . . . I. lilianae sp. n.
Ichthyocladius neotropicus Fittkau
Ichthyocladius neotropicus Fittkau, 1974: 101.
Material examined
Paratypes: Peru, Rio Tambopata, tributary of upper RioMadeira, one male pupa, one pharate female pupa, one larva,5–6.XI.1951, H. Heller (ZSM).
The species is described in detail by Fittkau (1974).However, his adult specimens were pharate and the illustrated
pharate male appears to be lost. Among other things it thusis not clear how many flagellomeres are present. Fittkau(1974) mentions that the male antenna is as in the female,which could mean that there are six flagellomeres only. Tem-porals also are mentioned to be absent. However, the tempo-ral setae present in the new species described here are tinyand could easily be overlooked.
Fittkau (1974) also mentions that the claws of the anteriorparapods are pale, thus contrasting with the black claws ofthe posterior parapods. In the examined larvae, however, theclaws of the anterior parapods are brown although not as darkas in the new species described below.
Fittkau overlooked the anal macrosetae of the pupae. Inaddition to three ventromedian, strongly curved, slendermacrosetae, slightly widened in apical third, there is oneminute apical seta as also observed in most other Ichtyocla-dius exuviae.
Ichthyocladius kronichticola sp. n. (Figs. 1–20)
Type material
Holotype male with larval and pupal exuviae: Brazil, SãoPaulo State, Parque Estadual Intevales, Iporanga, Rio daMortes, 17.VIII.2000, H.F. Mendes (MZUSP). Paratypes:one male, Brazil, São Paulo State, Iporanga, Petar,24°33¢52≤S 48°40¢35≤W, 3.IV.1997, S. Buck (ZSM); fivelarvae, São Paulo State, Parque Estadual Intervales, Ipo-ranga, Poços Altos, 1.II.2000, P.C. Bispo, A.S. Melo & V.R.de Ribeiro (MZUSP, ZMBN, ZSM).
Etymology
From Kronichthys, the genus of Loricariid fishes on whichthe larvae where situated, and – cola, Latin suffix meaninginhabiting. Regarded as a noun in apposition.
Description
Male imago (n = 2, except when otherwise stated). Total length 2.02–2.30mm. Wing length 1.32–1.34mm long.Total length/wing length 1.51–1.80. Wing length/length ofprofemur 3.29–3.52. Thorax and head dark brown; abdomendark brown with posterior 1/4 of segment VI and posteriorhalf of VII pale; coxa, trochanter, femora and tibia darkbrown with setae on fore tibia in pale spots, tarsi pale; wingsdark.
Head (Fig. 1). Antenna (Fig. 3) with 12 flagellomeres, AR0.21–0.28. Ultimate flagellomere 100–136 mm long. Tempo-ral setae consisting of three minute outer verticals and aboutfive minute inner verticals. Clypeus with 9–13 setae. Tento-rium (Fig. 2) 135 mm long, 38 wide. Palpomere 43–45 mmlong.
Thorax (Fig. 4). Dorsocentrals 14–19, in 1–3 rows; pre-alars 4–7. Scutellum with nine setae.
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New Species of Ichthyocladius 19
Wing (Fig. 8). VR 1.18–1.20. Clavus length 735–776mmlong. Clavus length/wing length 0.55–0.56. Cu/Wing length0.61 (1).
Legs (Table 1). Spur of front tibia (Fig. 5) 21 mm long;spurs of mid tibia (Fig. 6) 21–23 and 37–41mm long; spursof hind tibia (Fig. 7) 18 and 21(1) mm long. No comb setae.Pseudospurs each about 10 mm long.
Hypopygium (Figs. 9, 10). Tergite IX with 14–17 apical dorsal setae and 12–14 apicoventral setae. Phal-lapodeme 60 mm long. Gonocoxite 184mm long, inferiorvolsella moderately developed; gonostylus 82 mm long, with a few weak setae at apex, longest 5–7 mm long. HR 2.25;HV 2.46–2.80.
Figs. 1–10. Ichthyocladius kronichticola sp. n., male imago: (1) Head. (2) Tentorium and cibarial pump. (3) Antennal apex. (4) Thorax. (5)Apex of front tibia. (6) Apex of mid tibia. (7) Apex of hind tibia. (8) Wing. (9) Tergite IX and dorsal aspect of gonocoxite and gonostylus.(10) Hypopygium with tergite IX and gonostylus removed: left dorsal aspect, right ventral aspect.
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20 H.F. Mendes et al.
Figs. 11–20. Ichthyocladius kronichticola sp. n., pupa (Figs. 11–15) and larva (Figs. 16–20). (11) Tergites. (12) Sternites. (13) Frontalapotome and antennal sheath. (14) Thorax and wing sheath. (15) Macrosetae. (16) Labrum. (17) Mentum. (18) Mandible. (19) Antenna. (20)Apex of abdomen.
Table 1. Lengths (in mm, n = 1–2) and proportions of the legs of Ichthyocladius kronichticola sp. n.
fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR
P1 380–400 360–380 300–312 120 64–68 32 32–40 0.79–0.86 4.14–4.25 2.44–2.53 2.5–2.7P2 360 440–460 260–280 128 72 32 40 0.56–0.64 3.97 2.86–3.16 1.9P3 528 508 328 144 80 32 28 0.65 4.80 3.16 2.4
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New Species of Ichthyocladius 21
Pupa (n = 1). Total length 2.79mm. Exuviae dark amber withrows of very dark hooklets, genital sacs darker, abdominalsegments darker in median and lateral parts.
Cephalothorax. Frontal apotome and antennal sheath as inFig. 13. Thoracic setae (Fig. 14) less than 50 mm long,Dc1–Dc2 34mm, Dc2–Dc3 60mm, Dc3–Dc4 69mm.
Abdomen (Figs. 11, 12). Tergite (T) II–VII with strongmedian shagreen spinules placed on protuberance, T VIII andIX with strong median shagreen, genital sac apically wrin-kled and with a few minute spinules. Sternites (S) I and IIwith weak shagreen, S III–VIII with strong shagreen. Sha-green on sternites weaker than on tergites. T I with 33strongly reduced caudal hooklets or spines of which two aredistinct, dark hooklets; number of subequal, stronger hook-lets on T II–V as 41, 46, 37, 33; on S IV–VII as 42, 39, 38,32. Macrosetae (Fig. 15) scimitar-like, slender in basal 2/3wide from apical third, tapering to sharp point, about 45 mmlong. Additional hair-like seta placed slightly ventral of apexabout 19mm long.
Fourth instar larva (n = 3–4, except when otherwise stated).Total length 3.0–3.8mm. Head capsule 0.32–0.34mm long.Postmentum 126–135 mm long. Head light brown, bodygreyish black.
Head. Antenna as in Fig. 19, antennal segments (in mm):26–30, 9–11, 8–9, 4–8. AR 1.08–1.33. Basal segment 15–17,16mm wide; ring organ 21 mm (1) from base; blade 24mm (1)long; accessory blade not measurable. Labrum as in Fig. 16.Premandible 50–52 mm (2) long. Mandible (Fig. 18) 79–90mm long. Mentum (Fig. 17) 69–83, 76 mm wide; with16–18 median and five pairs of lateral teeth; distance betweensetae submenti 36–45, 39 mm; ventromental plate 9–15, 12mm wide.
Body. Some claws of anterior parapod with a few weak teeth. All setae on thorax and abdominal segmentsI–VIII less than 15mm long. Procercus (Fig. 20) 4–8 mm (2) high; 8–9 mm wide; with 4 anal setae 49 mm (2) long. Anal tubules 34 mm (1) long, 23 mm wide at base. Posteriorparapods with 13–15 black claws, including 1–3 smallerclaws.
Third instar larva (n = 1–2). Total length 2.22mm. Headcapsule 0.23–0.24mm long. Postmentum 90–105 mm long.Head light brown, body greyish black.
Head. Antennal segments (in mm): 15–17, 8, 8–9, 4. AR0.73–0.82. Basal segment 11–12mm wide, ring organ 11 mmfrom base, blade 19–21 mm long, accessory blade 9 mm long.Premandible 33mm (2) long. Mandible 60–64 mm long.Mentum 26–31 mm wide, with 18 median and five pairs oflateral teeth, distance between setae submenti 26–31 mm,ventromental plate 6–8 mm wide.
Body. All setae on thorax and abdominal segments I–VIIIless than 15 mm long. Procercus 6 mm high, 8mm wide, withfour anal setae 45mm long. Posterior parapod 94 mm long.Anal tubules 23 mm long, 11mm wide at base.
Diagnostic characters
The male imago of I. kronichticola sp. n. differs from that ofI. neotropicus by having a larger inferior volsella, presenceof outer verticals and 12 flagellomeres. The pupa differs fromother species by having strong median spinules placed on aprotuberance on tergites II–VII. The larva has a relativelylarger seta subdentalis, and a small, but distinct procercuswith four anal setae.
Biology
The larvae were taken on Kronichthys sp. Most of the hostfishes were collected under decayed leaves along the banksof a fourth order stream, where the water flowed very slowly(Melo & Froehlich, 2001, stream number 9). The fishes weresampled at 9 to 10 a.m., when they were inactive. A total of59 specimens were collected, but only five were carryingIchthyocladius larvae. Some other Loricariidae specimenswere also collected, three specimens of Neoplecostomus sp.and 12 specimens of Harttia spp., but all without Ichthy-ocladius larvae.
The fishes were brought to the laboratory individually inplastic bags. In order to rear Ichthyocladius adults associatedwith the immatures, each fish carrying only one larva wasplaced in a 15-litre aquarium. The only successful rearingwas obtained two days after the fishes were collected.Another adult was reared by the same method, but theexuviae was unfortunately not found.
Ichthyocladius lilianae sp. n. (Figs. 21–31)
Type material
Holotype: pharate male pupa with associated larva: Brazil, Minas Gerais, Rio Sao Francisco, 20°30¢0≤S46°50¢0≤W, 31.XII.1995, L. Casatti (MZUSP). Paratypes: 13larvae, same data as holotype except 1993–2000 (MZUSP,ZMBN).
Etymology
Named after Lilian Casatti, who collected the material.
Description
Male imago (n = 1).Head. Antenna with 12 flagellomeres, AR 0.28. Ultimate
flagellomere 116 mm long. Temporal setae consisting of at least one minute outer vertical and one minute inner vertical. Clypeus with 13 setae. Palpomere 43–45mm long.
Legs (Table 2). Spur of front tibia 19 mm long; spurs ofhind tibia 19 and 34mm long. No comb setae.
Hypopygium (Figs. 21, 22). Tergite IX with altogether 25setae. Phallapodeme 105 mm long, apodeme lobe prominent.
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22 H.F. Mendes et al.
Gonocoxite 176 mm long, inferior volsella moderately devel-oped; gonostylus 98mm long. HR 1.81.
Other details not measurable.
Pupa (n = 1). Total length 2.79mm. Exuviae pale withmargins of wing sheaths yellowish brown and hooklets yellowish.
Abdomen (Figs. 23, 24). Tergites (T) II and III with weakand sparse shagreen, T IV with slightly stronger and moreextensive shagreen, T V–IX with strong and extensive sha-green, genital sac mostly smooth. Sternites (S) I with weakand sparse shagreen, S II–VIII with stronger and more exten-sive shagreen. T I with 33 strongly reduced caudal hookletswidely interrupted medially; number of subequal, strongerhooklets on T II–V as 36, 37, 34, 36; on S IV–VII as 29, 33,29, 26. Macrosetae (Fig. 25) about 34 mm long, scimitar-like,broad near base to slightly past middle, then tapering to sharppoint; positioned at apex. Additional minute apical setaapparently present lateral of macrosetae.
Fourth instar larva (n = 8–11, except when otherwisestated). Total length 1.7–3.0, 2.4mm. Head capsule0.32–0.36, 0.34mm long. Postmentum 131–165, 145mmlong. Head light brown with postoccipital margin, postero-
lateral parts of gula, mentum, mandible and premandibledark brown.
Head. Antenna as in Fig. 30. Length of antennal segments(in mm): 19–24, 21; 6–9, 7; 5–7, 6; 2–5, 4. AR 1.13–1.43,1.27. Basal segment 9–13, 11 mm wide; ring organ 11–18, 14mm from base; basal setal mark 5mm from base, distal mark17–19, 18mm (6) from base; blade 12–15, 13 mm long; acces-sory blade 6–7, 6 mm (5) long. Labrum as in Fig. 26. Premandible (Fig. 27) 50–59, 59 mm long; with three darkinner teeth followed by three broadly lamellate fused teeth,apparently with vestigial brush. Mandible (Fig. 29) 76–104,84mm long. Mentum (Fig. 28) 73–85, 78 mm wide; with 20median and five pairs of lateral teeth; distance between setae submenti 33–45, 41 mm; ventromental plate 5–8, 7 mm wide.
Body. Some claws of anterior parapod with a few weakteeth. All setae on thorax and abdominal segments I–VIII lessthan 15 mm long. Procercus (Fig. 31) 5–7, 6 mm (2) high;9–13, 11mm wide; with four anal setae 33–59, 41 mm long.Anal tubules 52–71 mm (3) long, 19–24mm (3) wide at base.Posterior parapods with 12 black claws.
Third instar larva (n = 4–5, except when otherwise stated).Total length 1.53–1.88, 1.69mm. Head capsule 0.23–0.25,
Figs. 21, 22. Ichthyocladius lilianae sp. n., male imago. (21) Tergite IX and dorsal aspect of gonostylus. (22) Hypopygium with tergite IXand gonostylus removed: left dorsal aspect, right ventral aspect.
Table 2. Lengths (in mm, n = 1) and proportions (approximate) of the legs of Ichthyocladiuslilianae sp. n.
fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR
P1 431 420 311 109 75 41 43 0.74 4.34 2.94 –P2 371 334 278 105 64 36 41 0.83 4.00 2.54 –P3 – – – 120 79 30 43 – – – –
NNFE391015 8/17/04 2:14 PM Page 22
New Species of Ichthyocladius 23
Figs. 23–31. Ichthyocladius lilianae sp. n., pupa (Figs. 23–25) and larva (Figs. 26–31). (23) Tergites. (24) Sternites. (25) Macrosetae. (26)Labrum. (27) Premandible. (28) Mentum. (29) Mandible. (30) Antenna. (31) Apex of abdomen.
0.24mm long. Postmentum 90–113, 103mm long. Head lightbrown, body greyish black.
Head. Antennal segments (in mm): 13–17, 15; 6–7, 6; 2–6,5; 2–4, 4. AR 0.86–1.17, 0.99. Basal segment 7–8, 8mm wide;ring organ 7–12, 11mm from base; blade 9–12, 11 mm long;accessory blade 6mm (2) long. Mandible 59–73, 65 mm long.Mentum 40–52, 49mm wide; distance between setae submenti24–36, 29mm; ventromental plate 4–5, 5 mm wide.
Body. Procercus 6–7, 7 mm high; 7–9, 8mm wide; withfour anal setae 38–50, 44 mm long. Posterior parapod
119–152, 139mm long. Anal tubules 21–28, 24mm long;11–14, 12mm wide at base.
Diagnostic characters
The male imago of I. lilianae sp. n. differs from that of I.kronichticola sp. n. by having a smaller inferior volsella, and from I. neotropicus by having more numerous flagel-lomeres and only two temporals. The males of I. lilianaeand I. neotropicus, however, are known only from mature
NNFE391015 8/17/04 2:14 PM Page 23
24 H.F. Mendes et al.
pupae. The pupa of I. lilianae differs from that of I. kronichticola by lacking strong median shagreen spinulesplaced on protuberances on tergites II–VII, from I. neotrop-icus by having the caudal hooklets of tergite II as strong asthose on more posterior tergites, and from the unnamedspecies by having pale exuviae and strongly curved, scimi-tar-like anal macrosetae. The larva has a relatively larger setasubdentalis, and a small, but distinct procercus with four analsetae.
Biology
The larvae were collected on Hypostomus cf. garmani(Regan, 1904) and Harttia sp.
Ichthyocladius sp. Rio Marauiá (Figs. 32, 33)
Material examined
Brazil, Amazonas State, Rio Marauiá, upper half fromCachoeira S. Antônio, surface drift (Abschaum), one femalepupal exuviae, 22.I.1963, E.J. Fittkau (ZSM).
Description
Pupa (n = 1). Total length 2.93mm. Exuviae dark amber withvery dark rows of hooklets, abdominal segments VII and VIIIdarker laterally.
Cephalothorax. All thoracic setae less than 50 mm long,Distance Dc1–Dc2 19mm, Dc2–Dc3 71mm, Dc3–Dc4 81mm.
Abdomen (Fig. 30). Abdominal apex rounded, graduallynarrowed between segment VIII and IX. Tergite (T) II–VIIwith median shagreen spinules not on protuberance. T I with16 weak and small medial hooklets or spines, none dark;number of caudal, subequal, stronger hooklets on T II–V as31, 40, 38, 34; on S IV–VII as 30, 23, 32, 26. Macrosetae asin Fig. 33.
Ichthyocladius sp. Argentina (Figs. 34, 35)
Material examined
Argentina, Parque Nacional Iguazú, river before the falls,three male and one female pupal exuvia, 4.XII.1996, F. Reiss(ZSM).
Description
Pupa (n = 4). Total length 2.65–3.12, 2.79mm. Exuviae darkamber with very dark rows of hooklets, genital sacs darkerlaterally.
Cephalothorax. All thoracic setae less than 50 mm long,Distance Dc1–Dc2 41–94, 77mm; Dc2–Dc3 8–60, 33mm;Dc3–Dc4 124–150, 137mm.
Abdomen (Figs. 32). Abdomen abruptly narrowedbetween segment VIII and IX. Tergite (T) II–VII with medianshagreen spinules not on protuberance. T I with 27–45, 37weak and small medial hooklets; number of caudal, subequal,stronger hooklets on T II–V as 34–45, 39; 30–39, 34; 27–43,
Figs. 32–35. Ichthyocladius sp. Rio Marauiá (Figs. 32, 33) and Ichthyocladius sp. Argentina (Figs. 34, 35). (32 & 34) Tergites. (33 & 35)Macrosetae.
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New Species of Ichthyocladius 25
Table 3. Characters and states used in the phylogenetic analyses.
Imagines1. Antennal ratio: (0) higher than 0.9; (1) 0.46–0.9; (2) lower than 0.46.2. Number of flagellomeres: (0) 13; (1) fewer.3. Male antennal apex: (0) without subapical seta; (1) with.4. Female antennal apex: (0) without subapical seta; (1) with.5. Eyes: (0) bare, at most partly pubescent; (1) hairy or pubescent.6. Palpomeres: (0) five palpomeres of normal length; (1) four or less palpomeres or palpomeres strongly reduced in length.7. Sensilla clavata of palpomeres: (0) palpomere 3 at most with few sensilla in one group, palpomere 4 without sensilla; (1) in most
species at least female with sensilla in more than one group on palpomere 3 and palpomere 4 usually also with at least sensillaclavata, or, in Chaetocladius, numerous sensilla clavata at well developed sensillum coeloconicum.
8. Dorsomedian eye elongation: (0) moderately to well developed; (1) absent or very weak.9. Temporals: (0) inner verticals present or replaced by frontals, usually more outer verticals; (1) inner verticals absent, outer verticals
usually few.10. Antepronotal lobes: (0) broad, collar like, at most slightly narrowed medially; (1) distinctly narrowed medially.11. Dorsal antepronotals: (0) absent; (1) present.12. Humeral pit: (0) inconspicuous; (1) consisting of several smaller areas; (2) conspicuous, oval.13. Dorsocentrals: (0) erect; (1) decumbent.14. Dorsocentrals: (0) uniserial anterior; (1) bi–multiserial anterior.15. Dorsocentrals: (0) uniserial posterior; (1) bi–multiserial posterior.16. Acrostichals: (0) moderately long to long and strong, (1) short, or absent.17. Acrostichals: (0) starting in front; (1) starting some distance from antepronotum; (2) in centre of scutum. – Absence scored as (?).18. Acrostichals: (0) simple or absent; (1) often scalpellate.19. Prealars: (0) 1–5; (1) 6 or more.20. Supraalar(s): (0) present; (1) absent.21. Setae of preepisternum and/or anepisternum: (0) present; (1) absent.22. Scutellars: (0) uniserial; (1) bi–multiserial.23. Postnotum: (0) bare; (1) sometimes with setae. – Polymorphies are scored as synapomorphies, i.e. the tendency to have setae is
regarded as the synapomorphy. Otherwise the character would be uninformative since only some species of the included genera havesetae on the postnotum. To score the polymorphies would be the same as deleting the character.
24. Wing spots: (0) absent; (1) present.25. Wing membrane: (0) with setae; (1) bare.26. Wing membrane: (0) not to moderately punctated; (1) coarsely punctated.27. Anal lobe: (0) relatively well developed, often protruding; (1) weak, reduced or cuneiform wing.28. Costa: (0) distinctly extended; (1) moderately to not extended.29. R4+5: (0) ends above or distal to apex of M3+4; (1) ends proximal to apex of M3+4.30. Clavus: (0) absent; (1) present and ending in distal half of wing; (2) present and ending in proximal half of wing.31. Cu1: (0) not sinuous; (1) slightly sinuous; (2) strongly sinuous.32. VR: (0) <1.06; (1) 1.06–1.40; (2) >1.40.33. Anal veins: (0) An1 extending well beyond cubital fork and An2 conspicuous; (1) anal veins shorter.34. R veins: (0) setae present on R, R1 and usually R4+5 in both sexes; (1) setae present on R, absent on R1 and often R4+5 of male, at most
absent on R1 in female; (2) setae absent on R of male, present in female; (3) setae absent on R and R1 of both sexes, at most 1 apicalseta on R4+5.
35. Squama: (0) with setae; (1) bare.36. Leg ratio of male (LR1): (0) higher than 0.8; (1) 0.5–0.8; (2) lower than 0.5.37. Trochanter: (0) without keel; (1) at least fore trochanter with keel.38. Ta5: (0) not cordiform; (1) cordiform.39. Tibial spurs: (0) with distinct lateral denticles; (1) denticles indistinct or absent.40. Inner tibial spur of hind leg: (0) At least 1/2 as long as outer spur; (1) shorter; (2) absent with also second spur of mid leg absent.41. Hind tibial comb: (0) well developed, occupying full width of tibia; (1) weak or absent.42. Hind tibial comb: (0) with less than 13 setae; (1) often conspicuous with 13 or more setae of which some about as long as longest
spur.43. Pseudospurs: (0) present; (1) absent.44. Sensilla chaeticae of tarsi: (0) present; (1) absent.45. Pulvilli: (0) present and distinct; (1) absent or vestigial, less than 1/2 claw length.46. Anal point: (0) absent or small and anterior on tergite; (1) represented by hump-like extension of tergite or if absent represented by
some stronger median setae; (2) well set off and posterior on tergite.47. Anal point: (0) not with spatulate microtrichiae-free apex; (1) often with small or large spatulate, microtrichiae-free apex.48. Anal point: (0) not very broad and rounded to bluntly triangular; (1) often conspicuously broad and rounded to bluntly triangular.49. Setae on anal point or posterior on tergite IX: (0) conspicuous, bristle-like to lamellate; (1) moderately developed; (2) short, weak or
absent.50. Tergite IX: (0) normal, not very large; (1) large, covering most of hypopygium.51. Phallapodeme: (0) not pointed and recurved; (1) pointed and recurved.
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26 H.F. Mendes et al.
Table 3. Continued
52. Superior volsella: (0) present; (1) absent.53. Superior volsella: (0) not bare and plate-like; (1) bare and plate-like.54. Gonostylus: (0) simple; (1) double.55. Heel of gonostylus: (0) absent; (1) present at least in many species.56. Transverse sternapodeme: (0) convex; (1) straight or concave; (2) absent, sternapodeme triangular.57. Oral projections of transverse sternapodeme: (0) strongly developed; (1) weak to moderately developed; (2) absent.58. Crista dorsalis: (0) evident, triangular or rounded preapical; (1) elongate, low; (2) not evident/weak.59. Megaseta: (0) present; (1) absent.60. Virga: (0) consisting of tight cluster of at least six spines or two groups of very strong spines; (1) virga not consisting of cluster or
groups of spines.61. Virga: (0) virga when present consisting of cluster or field of spines or few spines with lateral lamellae; (1) consisting of 1–5 short
partly fused, sometimes plate-like spines without distinct lateral lamellae.62. Virga: (0) virga when present consisting of cluster or field of spines or few spines without lateral lamellae; (1) consisting of 1–3 long,
median, usually fused spines and distinct lateral lamellae.63. Gonocoxapodeme: (0) absent, short and straight or evenly curved and ending at base of gonapophysis; (1) continuing basal of vagina
or at least past base of gonapophysis.64. Female tergite IX: (0) undivided; (1) divided by caudal concavity or notch; (2) divided into two setigerous protrusions.65. Female gonocoxite IX: (0) large, projecting; (1) moderately developed to reduced.66. Female gonocoxite IX: (0) with several long setae; (1) with less than three long setae; (2) with some long and some short setae; (3)
with short setae only.67. Gonapophysis VIII: (0) undivided; (1) divided with ventrolateral lobe much smaller and more or less brush-like, or with lobes of
about same size; (2) divided with dorsomesal lobe smaller and with anterior rounded projection; (3) divided with dorsomesal lobenarrow, often line-like.
68. Apodeme lobe: (0) not apparent; (1) well developed, but not meeting at mid line and with microtrichiae; (2) meeting at mid lineand/or with microtrichiae; (3) fully sclerotized.
69. Number of seminal capsules: (0) 3; (1) 2.70. Seminal capsules: (0) spherical to ovoid, small or of normal size; (1) large, spherical to elongate ovoid.71. Seminal capsules: (0) at least partly coloured; (1) often completely pale. – Polymorphies are scored as synapomorphies as no genera
have all included species with pale capsules and the character otherwise would be uninformative.72. Opening of spermathecal ducts: (0) separate; (1) common.73. Spermathecal ducts: (0) not fused; (1) partly fused ducts before common opening.74. Bulbs of spermathecal ducts: (0) absent; (1) present.75. Spermathecal ducts: (0) straight; (1) with bend or loop.
Pupa76. Frontal apotome: (0) without warts or tubercles; (1) with warts or tubercles.77. Frontal setae: (0) present; (1) absent.78. Thoracic horn: (0) present; (1) absent.79. Thoracic horn: (0) not rounded to ovoid; (1) often rounded to elongate ovoid.80. Thoracic horn: (0) not with bulbous base and narrow apical portion; (1) with. – Eukiefferiella is scored as (1) since most species
have a thoracic horn with bulbous base and species without thoracic horn apparently have this secondarily reduced.81. Thorax: (0) mostly smooth to wrinkled; (1) mostly tuberculose or spinulose.82. Thorax: (0) without tubercle(s) medially or in front; (1) with. – The polymorphy for Cardiocladius s. lat. is scored as a
synapomorphy as otherwise the presence of a tubercle will appear as an autapomorphy for Ichthyocladius.83. Wing sheath: (0) without pearls; (1) with.84. Dorsocentrals: (0) anterior two and posterior two (three) paired, anterior three grouped, all in row or 2–3 dorsocentrals only; (1)
posterior three grouped or all four together.85. Tergites II–VIII: (0) without posterior spine, or tubercle row(s), but may have caudal hooklets; (1) some with spines or tubercles.86. Median field of tergite IV: (0) without discrete spine patch(es) or row(s); (1) sometimes with.87. Tergite I: (0) without posterior spine row(s); (1) with.88. Sternites II–VII: (0) without posterior spine row(s), but may have caudal hooklets; (1) some with spines or tubercles.89. Sternites or sternal conjunctives: (0) without caudal hooklets; (1) at least one with.90. Male sternite VIII: (0) without posterior spine or tubercle row(s); (1) with.91. Sternite II or II and III: (0) without anterior or median spine group; (1) with.92. Tergites and sternites: (0) with single or no posterior row of spines; (1) at least some with double to multiple row of spines.93. Tergite III: (0) without caudal hooklets; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets, which may be comb-
shaped. – Cricotopus absurdus type without thoracic horn and with pearl row, another type with hooklets on T III and PSB on II andIII. Characters 93–97 are likely to be interdependent and the presence of hooklets thus is receiving higher weight than othercharacters. However, several other characters also are likely to be genetically connected and the hooklets certainly are of significance.
94. Tergite IV: (0) without caudal hooklets; (1) with minute caudal hooklets; (2) with conspicuous caudal hooklets, which may be comb-shaped.
95. Tergite V: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minutecaudal hooklets; (2) with conspicuous caudal hooklets which may be comb-shaped.
NNFE391015 8/17/04 2:14 PM Page 26
New Species of Ichthyocladius 27
Table 3. Continued
96. Tergite VI: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minutecaudal hooklets; (2) with conspicuous caudal hooklets.
97. Tergite VII: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-like anteriorly directed; (1) with minuteor conspicuous caudal hooklets. – Polymorphies are scored as synapomorphies as the character otherwise would be uninformative(See character 23).
98. Caudal hooklets of tergite II: (0) present, (1) absent.99. Pedes spurii B: (0) present; (1) absent.
100. Pedes spurii A on sternite IV: (0) present; (1) absent.101. Pedes spurii A on sternite VI: (0) present; (1) absent.102. Tergal conjunctives or terga posterior of tergal spines: (0) without spinules, but may have hooklets in single row; (1) with spinules
which may be hooklet-like recurved.103. Spinules on tergal conjunctives: (0) absent or not hooklet-like recurved or anteriorly directed; (1) hooklet-like recurved or anteriorly
directed spinules in about three rows.104. Taeniate L setae: (0) present; (1) absent.105. Anal lobe: (0) not extended into projections; (1) extended posteriorly into cylindrical projections with macrosetae at apex.106. Anal lobe: (0) not with apical projections or extensions; (1) at least sometimes with apical spurs or extended distal of macrosetae.107. Apical spines of anal lobe: (0) absent; (1) present.108. Anal lobe: (0) with fringe of setae; (1) without fringe of setae.109. Inner margin of anal lobe: (0) without long seta; (1) with.110. Anal macrosetae: (0) subequal; (1) differing; (2) absent or one only.111. Anal macrosetae: (0) not conspicuously curved and scimitar-like; (1) sometimes conspicuously curved and scimitar-like.112. Anal macrosetae: (0) not short and spine-like, but may be short and hair-like or absent; (1) short and spine- or thorn-like; (2) absent.113. Width of anal macrosetae or apical spines: (0) less than 5 mm; (1) 5 mm or more.114. No. of anal macrosetae: (0) three or more; (1) 2; (2) 0–1.
Larva115. Antenna: (0) with 6–7 segments; (1) with five; (2) with 3–4.116. Ultimate antennal segment: (0) normal; (1) whip- or thread-like.117. Antenna: (0) reduced, less than half mandible length; (1) 1/3 as long as head capsule or shorter, but not reduced; (2) longer.118. Second antennal segment: (0) undivided, fully sclerotised; (1) divided or partly unsclerotised.119. Lauterborn organs: (0) moderately large to well developed; (1) weak or absent.120. Antennal blade: (0) short to moderate length, shorter than flagellum when antenna of normal length; (1) conspicuous, longer than
flagellum except when flagellum extremely long.121. S I: (0) plumose, branched, pectinate or palmate; (1) bifid or simple.122. S I: (0) not bifid; (1) bifid.123. S I: (0) not simple; (1) simple.124. Labral lamella: (0) with pectinate, plumose or rugose apex, mostly well developed; (1) weak, no apical teeth or plumosity; (2) absent.125. Chaetulae laterales: (0) simple or reduced; (1) at least one serrated or plumose.126. Premandible: (0) simple; (1) with two or more teeth.127. Premandibular brush: (0) present; (1) absent.128. Mandible: (0) with 2–3 inner teeth; (1) with four or more.129. Mola of mandible: (0) smooth; (1) with teeth or spines.130. Seta interna of mandible: (0) with smooth, slightly plumose laterally or apically, or serrate branches; (1) branches conspicuously
branched; (2) seta interna absent.131. Median tooth of mentum: (0) single; (1) double, bifid or with several median teeth.132. Mentum: (0) median group of 4–20 teeth not forming straight line on strongly curved mentum; (1) median group of 4–20 teeth
forming straight line on strongly curved mentum so that lateral teeth not are visible on uncompressed larvae.133. Teeth of mentum: (0) 15 or more teeth; (1) 11–14 teeth; (2) fewer.134. Lateral teeth of mentum: (0) outer tooth not larger or higher than one of the inner teeth; (1) clearly larger or higher.135. Ventromental plates: (0) well developed, extending past lateral teeth on flattened mentum; (1) reduced or weak.136. Ventromental plates: (0) without setae (beard) underneath; (1) with setae underneath.137. Setae submenti: (0) situated at level of base of outer lateral tooth or higher; (1) lower.138. Claws of anterior parapods: (0) with relatively distinct teeth; (1) smooth or teeth very indistinct.139. Setal tufts on abdomen: (0) absent; (1) often present.140. Procercus: (0) well developed; (1) reduced or absent.141. Anal setae: (0) five or more setae, none conspicuously long; (1) 3–4 setae, none conspicuously long; (2) 0–2 not conspicuously long
anal setae; (3) two or more setae with one or two conspicuously long.142. Supraanal setae: (0) weak, shorter than 1/2 length of anal setae when these well developed or shorter than 2/3 when anal setae short;
(1) well developed, longer than 1/2 length of anal setae when anal setae long, longer than 2/3 when anal setae short.143. Posterior parapods: (0) well developed; (1) small, digitiform; (2) absent to weak but not digitiform.144. Anal tubules: (0) at least 1/2 length posterior parapods; (1) shorter than 1/2 length posterior parapods or these absent; (2)
conspicuously long and narrow.
NNFE391015 8/17/04 2:14 PM Page 27
28 H.F. Mendes et al.
Tabl
e 4.
Cha
ract
er s
tate
s fo
r ch
arac
ters
1–1
44 i
n so
me
gene
ra o
f O
rtho
clad
iina
e. P
olym
orph
ies:
A =
0&1,
B =
0&1&
2, C
=1&
2, D
=1&
2&3,
E =
0&2,
F =
0&3,
G =
2&3,
H =
0&1&
2&3,
I =
0&2&
3, J
=1&
3.
Cha
ract
er n
o.1
23
45
67
89
11
11
11
11
11
22
22
22
22
22
33
33
33
30
12
34
56
78
90
12
34
56
78
90
12
34
56
Ant
illo
clad
ius
Sæ
ther
A0
00
00
11
A0
00
00
01
11
00
10
00
A0
1A
00
01
11
A1
Bri
llia
Kie
ffer
00
11
00
01
00
10
0A
11
?0
10
11
00
01
01
00
01
00
01
Bry
opha
enoc
ladi
us0
00
A0
00
0A
00
00
0A
AA
00
A1
00
01
AA
A0
00
10
0A
AT
hien
eman
nC
ardi
ocla
dius
Kie
ffer
0
00
A0
00
10
0A
00
A0
10
?0
?1
10
01
10
10
0A
00
AA
1s.
l.
Cha
etoc
ladi
us K
ieff
erB
AA
0A
01
A0
00
20
AA
10
0A
A1
00
01
10
10
0A
00
AA
1C
oryn
oneu
raW
inne
rtz
BA
AA
10
0A
A0
0A
1A
A1
00
AA
AA
1A
1A
01
00
01
1B
0B
Cri
coto
pus
v.d.
Wul
pB
AA
A1
00
AA
00
A1
AA
10
0A
AA
A1
A1
A0
A0
00
10
B0
BD
iplo
clad
ius
Kie
ffer
00
00
10
01
11
00
00
01
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Boo
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20
31
2&
Sæ
ther
NNFE391015 8/17/04 2:14 PM Page 28
New Species of Ichthyocladius 29P
rops
iloc
erus
Kie
ffer
AA
00
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11
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Pse
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Tem
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Cha
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31
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0
11
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1D
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10
00
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10
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31
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11
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10
10
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10
10
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02
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00
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10
01
10
01
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0
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10
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10
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10
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10
11
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10
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01
00
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01
01
02
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31
10
01
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00
11
01
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20
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01
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20
10
01
20
20
01
00
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pärc
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ladi
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10
01
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21
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01
01
00
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01
11
00
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pärc
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00
11
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00
12
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01
00
10
12
00
00
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12
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11
01
Icht
hyoc
ladi
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01
01
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10
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22
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10
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10
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10
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00
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10
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Boo
thro
yd
NNFE391015 8/17/04 2:14 PM Page 29
30 H.F. Mendes et al.
Tabl
e 4.
Con
tinu
ed
Lim
noph
yes
Eat
on0
01
10
01
A1
10
0A
00
10
00
01
00
11
01
B1
23
11
00
1M
etri
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mus
v.d.
0
01
A0
10
11
20
01
00
10
00
0A
A0
A0
0A
AA
E3
11
A1
1W
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Oli
veir
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a0
01
10
01
01
00
02
00
10
00
01
00
10
0?
??
??
??
??
?W
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nbru
g &
Fi
ttka
uO
rtho
clad
ius
v.d.
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p0
01
00
0A
01
20
01
00
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00
01
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la0
01
10
11
11
20
11
00
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00
01
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10
10
21
21
11
00
1T
hien
eman
nPa
ram
etri
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mus
00
11
00
11
12
10
20
01
00
00
10
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00
12
12
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10
01
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11
11
00
11
12
00
10
01
00
00
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01
00
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& S
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01
01
01
01
00
AC
00
32
00
00
Pse
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clad
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0A
1B
01
00
0E
00
10
01
00
00
AE
01
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JC
1A
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01
10
01
10
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00
00
10
00
01
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10
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23
11
10
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01
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11
00
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21
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01
10
11
00
20
01
00
01
00
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10
00
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23
11
11
0T
hien
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H
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sch
Tem
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11
10
11
01
00
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11
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00
10
20
10
00
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& D
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aT
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11
10
11
11
00
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11
01
00
10
20
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31
10
1K
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00
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Cha
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77
77
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99
99
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11
11
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11
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45
67
89
01
23
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01
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45
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00
00
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34
56
78
Ant
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clad
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ther
01
10
11
00
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10
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01
NNFE391015 8/17/04 2:14 PM Page 30
New Species of Ichthyocladius 31D
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clad
ius
Kie
ffer
10
11
00
00
10
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00
10
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10
10
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11
Het
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00
00
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01
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01
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01
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10
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01
01
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01
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AA
00
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Pse
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01
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00
00
00
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11
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10
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dwar
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1
NNFE391015 8/17/04 2:14 PM Page 31
32 H.F. Mendes et al.
Tabl
e 4.
Con
tinu
ed
Cha
ract
er n
o.1
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
10
11
11
11
11
11
22
22
22
22
22
33
33
33
33
33
44
44
49
01
23
45
67
89
01
23
45
67
89
01
23
45
67
89
01
23
4
Ant
illo
clad
ius
Sæ
ther
00
01
1A
10
00
11
00
02
11
11
00
A0
21
10
11
01
20
01
Bri
llia
Kie
ffer
00
00
00
20
11
00
00
00
11
11
00
10
11
10
10
00
00
01
Bry
opha
enoc
ladi
us0
00
00
21
01
00
A1
01
20
11
00
2A
02
A1
01
10
12
02
1T
hien
eman
nC
ardi
ocla
dius
Kie
ffer
0
00
00
01
01
00
01
01
20
01
11
10
01
01
01
?0
1A
00
1s.
l.
Cha
etoc
ladi
us K
ieff
er0
A0
10
01
01
00
A0
00
AA
10
A0
1A
0C
0A
01
00
00
00
AC
oryn
oneu
raW
inne
rtz
10
00
00
20
20
10
10
12
11
01
0E
10
10
10
1?
00
10
00
Cri
coto
pus
v.d.
Wul
p0
A0
00
BC
0A
0A
01
AA
2A
A1
0A
00
01
01
01
01
00
00
AD
iplo
clad
ius
Kie
ffer
00
00
00
10
10
00
00
00
11
11
00
10
10
01
00
00
00
01
Dra
tnal
iaS
æth
er &
1
00
00
01
01
01
01
01
20
11
00
21
11
01
01
00
00
00
0H
alvo
rsen
Euk
ieffe
riel
la0
A0
00
EC
01
00
01
01
20
01
A1
0A
0C
01
01
A0
0A
A0
1T
hien
eman
nG
en.
n. n
ear
12
00
02
10
C0
00
00
02
00
01
00
10
11
10
10
00
10
01
Thi
enem
anni
ella
Gym
nom
etri
ocne
mus
0
00
00
C1
01
00
A1
01
20
11
00
21
02
A1
01
10
12
02
1G
oetg
hebu
erH
elen
iell
aG
owin
00
00
00
10
11
01
00
02
01
10
01
10
11
10
00
00
01
01
Het
erot
anyt
arsu
s0
00
00
02
02
00
10
00
11
11
00
01
01
00
00
00
00
A0
0S
pärc
kH
eter
otri
ssoc
ladi
us0
00
00
00
1C
01
A0
00
10
11
A0
1A
01
00
01
00
00
00
0S
pärc
kH
ydro
baen
usFr
ies
00
00
00
00
10
00
00
01
01
10
00
10
10
00
10
00
0A
00
Icht
hyoc
ladi
usFi
ttka
u?
A1
0A
02
0A
01
00
00
00
1A
A0
21
10
01
00
10
1A
00
1K
aniw
hani
wha
nus
02
02
02
20
00
00
11
0?
00
0A
00
00
20
11
1?
00
00
00
Boo
thro
ydL
imno
phye
sE
aton
00
00
00
10
10
1A
00
02
01
00
00
10
10
10
1A
00
01
0A
Met
rioc
nem
usv.
d.
0A
00
0B
10
10
AA
A0
AA
01
01
01
A0
10
10
11
00
01
0A
Wul
pO
live
irie
lla
00
01
10
??
??
??
??
??
??
??
??
??
??
??
??
??
??
??
Wie
denb
rug
&
Fitt
kau
Ort
hocl
adiu
sv.
d. W
ulp
00
00
0E
C0
10
00
11
02
AA
A0
0A
00
A0
1A
A0
10
00
0A
Para
kief
feri
ella
00
00
00
11
10
10
10
1C
0A
10
00
A0
10
00
0?
00
00
0A
Thi
enem
ann
Para
met
rioc
nem
us0
00
00
01
01
00
00
00
1A
11
00
A1
01
00
00
10
00
10
0G
oetg
hebu
erP
hyso
neur
a?
??
??
??
??
??
??
??
??
??
??
??
??
??
??
??
??
??
?Fe
rrin
gton
& S
æth
er
NNFE391015 8/17/04 2:14 PM Page 32
New Species of Ichthyocladius 33
34; 29–46, 35 on S IV–VII as 27–39, 33; 30–41, 34; 29–42,34; 26–39, 33. Macrosetae as in Fig. 35.
Male genital sac extending past anal segment by 38–83mm (3), female genital sac reaching apex of anal segment.
Phylogenetic analysis
A generic-level analysis based on the characters presented inTable 3 and the data matrix in Table 4 with DiplocladiusKieffer as outgroup was performed. In addition to Ichth-yocladius, all genera with caudal hooklets on tergites otherthan tergite II were included, i.e., genera near EukiefferiellaThienemann and Corynoneura. Presumed primitive generasuch as Brillia Kieffer, Diplocladius Kieffer, PropsilocerusKieffer, Heterotanytarsus Spärck, and Hydrobaenus Frieswere included in order to give direction of trends. Thesegenera also include genera with well developed, finely pecti-nate labral lamellae. Additional common, non-aberrantgenera representing different groups of genera also wereincluded. Oliveiriella Wiedenbrug & Fittkau according to thedescription (Wiedenbrug & Fittkau, 1997) has no supraalar,uniserial scutellars, the shorter hind tibial spur is more thanhalf as long as the outer, and pseudospurs and sensilla chaet-icae are not mentioned. However, M. Spies (personal com-munication) has re-examined the genus and found that asupraalar is present, scutellars are partly bi-serial, the shortertibial spur is less than half as long as the longer one, pseu-dospurs are absent and sensilla chaeticae present. The newgenus near Thienemanniella includes among others Thiene-manniella semifimbriata Sæther (Sæther, 1981: 32) and thistype is excluded from Thienemanniella for the charactersscored here. The characters are coded ordered to indicate thephylogenetic position of Ichthyocladius relative to knowngenera. The relationships among more basally placed generathus should not be taken for relevant.
Analyses were done with characters 30, 31, 34, 115, and124 ordered. When no characters were given weight, ninetrees were obtained each with a length of 588 steps, a con-sistency index (CI) of 0.30, a retention index (RI) of 0.46 anda rescaled consistency index (RC) of 0.14 (Fig. 36). Whenthe results were reweighted according to RC, a single treewith 590 steps after resetting to equal weights, CI-0.47, RI-0.70 and RC-0.33 was obtained (Fig. 37). TheCorynoneura group of genera appears monophyletic andincludes Ichthyocladius, while most genera of the Eukief-feriella group are included basally in the same monophyleticgroup. Dratnalia, Tvetenia and Oliveiriella Wiedenbrug etFittkau fall outside the group. When the results arereweighted according to the rescaled consistency indexTvetenia and Oliveiriella forms a separate group outside themain group by two nodes. The Bremer supports are shownabove each branch.
Several characters obviously are more important thanothers and should be weighted accordingly. Such weightingP
rops
iloc
erus
Kie
ffer
00
00
00
C0
10
10
00
00
11
11
00
10
A0
00
00
00
0A
00
Pse
ctro
clad
ius
Kie
ffer
0E
00
0E
10
10
10
00
02
00
10
0E
A0
10
01
01
00
A0
00
Pse
udor
thoc
ladi
us0
00
00
C1
01
0A
A0
00
20
01
00
11
02
A1
00
10
03
01
EG
oetg
hebu
erP
seud
osm
itti
a0
00
00
B2
00
01
11
10
21
10
A0
E0
0B
01
01
A0
1C
02
0E
dwar
dsR
heoc
rico
topu
s0
00
00
01
01
00
0A
A0
20
01
00
0A
01
00
10
00
00
A0
0T
hien
eman
n &
H
arni
sch
Tem
pisq
uito
neur
a0
20
20
21
01
01
11
10
21
00
10
10
01
01
01
00
01
00
0E
pler
& D
e la
Ros
aT
hien
eman
niel
la1
00
00
01
02
00
01
01
21
00
10
11
01
01
01
?0
0A
00
0K
ieff
erTo
kuna
gaia
Sæ
ther
0A
10
0E
10
10
10
10
12
A0
1A
00
10
10
10
10
00
00
0A
Tve
teni
aK
ieff
er1
00
00
01
01
00
A0
00
2A
01
01
0A
01
01
01
10
00
10
0
NNFE391015 8/17/04 2:14 PM Page 33
34 H.F. Mendes et al.
is rather subjective. However, with experience from relatedgroups and genera and giving weight to characters foundimportant as synapomorphies in other studies, characters 30,50, 54, 69, 80, 89, 94, 98, 99, 102, 104, 109, 135 and 136were given a weight of 10; characters 7, 11, 18, 31, 35, 45,
46, 83, 100, 101, 108, 122, 123, 127 a weight of 5. (Charac-ters 93 and 95–97 are not given extra weight since charac-ters 93–97 most likely are not genetically independent andthe presence of caudal hooklets is thus already weighted.)The resulting five trees obtained (Fig. 38) had 605–610 steps
Figs. 36–39. Parsimony analyses of the relationships between Ichthyocladius Fittkau and some selected Orthocladiinae. (36) Strict consen-sus tree of the 12 trees obtained with no characters weighted. (37) The single tree obtained with characters reweighted according to the rescaledconsistency index (RC). (38) Strict consensus tree of the five trees obtained with some characters weighted. (39) The single tree obtainedwhen this result reweighted according to RC.
NNFE391015 8/17/04 2:14 PM Page 34
New Species of Ichthyocladius 35
after resetting to equal weights, CI-0.35, RI-0.61 and RC-0.21. The Bremer supports above each branch are number ofsteps higher than 610 after resetting to equal weights.Reweighting according to RC resulted in a single tree of 599steps, CI-0.45, RI-0.73 and RC-0.36 (Fig. 39). The weightedresult has each of the Corynoneura and Eukiefferiella groupsseparately monophyletic before reweighting. The combinedgroup is monophyletic after reweighting with the Eukief-feriella group basal and paraphyletic.
The Bremer support is low when the characters areunweighted and not reweighted, high for the inclusion ofIchthyocladius in the Corynoneura group when the charac-ters are weighted or reweighted. The phylogenetic placementof Ichthyocladius appears to be confirmed, while the sug-gested relationship of the Eukiefferiella and Corynoneuragroup is strongly indicated.
Acknowledgements
We are very grateful to Marion Kotrba who provided andfacilitated the access to the Zoologische Staatssammlung col-lection of Chironomidae in Munich. The authors would liketo thank E.J. Fittkau and M. Spies for refereeing the paperand giving many helpful suggestions. The first author wassupported by the Brazilian Research Council of the SãoPaulo State (FAPESP proc. 00/05903-9 and proc 98/05073-4 Biota) while completing the paper.
References
Cranston PS, Oliver DR, Sæther OA (1983): The larvae ofOrthocladiinae (Diptera: Chironomidae) of the Holarcticregion – Keys and diagnoses. In: Wiederholm T, ed., Chi-ronomidae of the Holarctic region. Keys and diagnoses.Part 1. Larvae. Entomol Scand Suppl 19: 149–291.
Epler JH, De la Rosa CL (1995): Tempisquitoneura, a new genusof Neotropical Orthocladiinae (Diptera: Chironomidae)symphoretic on Corydalus (Megaloptera: Corydalidae). J N Am Benthol Soc 14: 50–60.
Fittkau JE (1974): Ichthyocladius n. gen., eine neotropischeGattung der Orthocladiinae (Chironomidae, Diptera) derenLarven epizoisch auf Welsen (Astroblepidae und Loricari-idae) leben. Ent Tidskr Suppl 95: 91–106.
Gonser T, Spies M (1997): Southern hemisphere Symbiocladius(Diptera, Chironomidae) and their mayfly hosts(Ephemeroptera, Leptophlebidae). In: Landolt P, Sartori M,eds., Ephemeroptera & Plecoptera: Biology – Ecology –Systematics, MTL – Fribourg, pp. 455–466.
Melo AS, Froehlich CG (2001): Macroinvertebrates in neotrop-ical streams: richness patterns along a catchment andassemblage structure between 2 seasons. J N Am BentholSoc 20: 1–16.
Sæther OA (1977): Female genitalia in Chironomidae and otherNematocera: morphology, phylogenies, keys. Bull Fish ResBoard Can 197: 1–211.
Sæther OA (1980): Glossary of chironomid morphology termi-nology (Diptera: Chironomidae). Entomol Scand Suppl 14:1–51.
Sæther OA (1981): Orthocladiinae (Diptera: Chironomidae)from the British West Indies, with descriptions of Antillo-cladius n. gen., Lipurometriocnemus n. gen. Compteros-mittia n. gen. and Diplosmittia n. gen. Entomol Scand Suppl16: 1–46.
Sæther OA (1986): The myth of objectivity – post-Hennigiandeviations. Cladistics 2: 1–13.
Sæther OA, Kristoffersen L (1996): Chironomids with ‘M-fork’.A reevaluation of the wing venation of the Corynoneuragroup (Insecta, Diptera, Chironomidae). Spixiana 19:229–232.
Svensson BS (1986): Eukiefferiella ancyla sp.n. (Diptera: Chi-ronomidae) a commensalistic midge on Ancylus fluviatilisMüller (Gastropoda: Ancylidae). Entomol Scand 17:291–298.
Wiedenbrug S, Fittkau EJ (1997): Oliveiriella almeidai(Oliveira, 1946), gen. nov., comb. nov., from South Americawith description of the pupae. Spixiana 20: 167–172.
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