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Morphology and Chaetotaxyof Larval Hydraenidae(Coleoptera) I: GenusLimnebius Leach, 1815 Basedon a Description of Limnebiuscordobanus d'OrchymontJuan A. Delgado a & Agustin G. Soler aa Departamento de Biologia Animal (Zoologfa),Facultad de Biologia , Universidad de Murcia ,Murcia, 30100, SpainPublished online: 30 Sep 2008.

To cite this article: Juan A. Delgado & Agustin G. Soler (1997) Morphology andChaetotaxy of Larval Hydraenidae (Coleoptera) I: Genus Limnebius Leach, 1815 Basedon a Description of Limnebius cordobanus d'Orchymont, Aquatic Insects: InternationalJournal of Freshwater Entomology, 19:1, 37-49, DOI: 10.1080/01650429709361634

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Aquatic Insects, Vol. 19 (1997), No. 1, pp. 37-49 0165-0424/97/1901-0037$12.00© Swets & Zeitlinger

Morphology and Chaetotaxy of Larval Hydraenidae(Coleoptera) I: Genus Limnebius Leach, 1815

Based on a Description of Limnebius cordobanusd'Orchymont

Juan A. DELGADO and Agustin G. SOLER

J.A. DELGADO and A.G. SOLER: Morphology and Chaetotaxy of Larval Hydraenidae(Coleoptera) I: Genus Limnebius Leach, 1815 Based on a Description of Limnebius cordo-banus d'Orchymont.

Aquatic Insects, Vol. 19 (1997) No. 1, pp. 37-49.

The morphology and chaetotaxy of the three larval instars of the genus Limnebius aredescribed and illustrated following the setal pattern of Limnebius cordobanus d'Orchymont,whose morphology is sufficiently representative of the genus. This is the first completestudy of larval chaetotaxy for a genus of Hydraenidae.

Keywords: Coleoptera, Hydraenidae, Limnebius, larvae, chaetotaxy.

J.A. DELGADO and A.G. SOLER, Departamento de Biologia Animal (Zoologfa), Facultadde Biologia, Universidad de Murcia, 30100 Murcia, Spain.

INTRODUCTION

The family Hydraenidae has been considered as a member of Hydrophiloidea,placed next to Hydrophilidae. However, authors who have studied larvae (Paulian,1941; Dybas, 1976) have consistently considered that Hydraenidae belong to theStaphylinoidea (Newton and Thayer, 1992). As in other groups, the biology andphylogeny of this family has been almost exclusively based on the observation ofadults (Perkins, 1980). Several workers have emphasized the importance of pre-imaginal stages to solve questions like these, but surprisingly few larvae havebeen described. Immature beetles provide an array of phylogenetic charactersthat supplement those wich can be observed at the adult stage (Wheeler, 1990),and many of such characters are related with the distribution of body setae andcampaniform sensilla.

Two principal systems of setal nomemclature have been proposed for namingsetae on larval Coleoptera: the Ashe-Watrous system (Ashe and Watrous, 1984)and the Bousquet-Goulet system (Bousquet and Goulet, 1984). These two sys-tems have been applied to different families of Coleoptera as Carabidae (Bous-quet and Goulet, 1984), Staphylinidae (Ashe and Watrous, 1984; Ashe, 1986),Dytiscidae (Wolfe and Roughley, 1985; Alarie, 1990; Alarie et al., 1990; Alarieand Harper, 1990), Leiodidae (Wheeler, 1990) or Histeridae (Kovarik and Pas-soa, 1993), but never in Hydraenidae.

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38 JUAN A. DELGADO AND AGUSTIN G. SOLER

B

Fig. 1. Limnebius cordobanus. Larval habitus of second instar; A: lateral view; B: dorsal view.Scale = 1 mm.

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LARVA OF UMNEBIUS (COL.: HYDRAENIDAE) 39

The purpose of this paper is to describe the morphology and chaetotaxy ofLimnebius and provide illustrations to facilitate their correct identification. Wehope that this work and the future chaetotaxal studies of other genera of Hydrae-nidae (Delgado and Soler, in prep.) might serve to clarify their natural historyand phylogenetic position.

MATERIAL AND METHODS

Material examined. The immature Limnebius used in this study were obtained by rearing adults in thelaboratory (ex ovo). Even though this method is complicated, the larvae that grow from well identifiedadults are the best material for chaetotaxal studies. Furthermore, obtaining sufficient material in nature(particularly the first instar) is difficult, especially in view of the minute size of the larvae. A secondproblem arises from the difficulty in correctly identifying the larvae and adults of the same species.However, some specimens of instars II and III identified ex societale imaginis were also studied. Theseimmatures were used exclusively as reference because of the above mentioned problem.

Several species of Limnebius from the south of the Iberian Peninsula were reared and their larvaeobtained. The biology of this genus and the rearing techniques used in this study will be described ina separate paper (Delgado and Soler, in prep.). Limnebius cordobanus d'Orchymont, 1938, waschosen to describe the setal pattern of Limnebius because this species has a representative or general-ized chaetotaxy and, more especially, because the larvae were available in reasonable quantities.When possible, at least six specimens were used to describe each character.

Larval study. The description of immature Limnebius is based on material cleared, glycerinatedand mounted on standard glass slides. The technique for studying larvae in glycerine has beendescribed by Bousquet and Goulet (1984). In those larvae which have a soft cuticle no clearingtechnique is necessary. For general studies under an optical microscope, the specimen was placed ona ringed slide filled with glycerine. The parts of the body to be examined under high magnificationwere detached and positioned on a slide with Hoyer's mounting medium and covered with a cover-slip. Drawings were prepared using a microscope with a drawing mirror.

Chaetotaxal analysis. The system for naming setae and pores used in this article closely followsthat of Ashe and Watrous (1984). In general, we have found that the setal patterns of Limnebius andother genera of Hydraenidae do not differ greatly from those found in Staphylinidae (Ashe, 1986) orLeiodidae (Wheeler, 1990). However, an important modification to the Ashe-Watrous system hasbeen proposed. Wheeler (1990) and Alarie et al. (1990) suggested that the value of chaetotaxy isenhanced when the primary setae are separated from the secondary setae which are added in laterinstars. This fact makes it easier to determine of serial homology among different segments. Weagree with these authors and took the first instar as the base for our chaetotaxal analysis.

The description is organized according to body region. Thorax and abdomen share a similar codewhereas the head has a different setal pattern and code. We agree with Ashe and Watrous (1984) inselecting the prothorax as the base for determining serial homology in contrast to Kovarik and Passoa(1993) who selected the first abdominal segment. In staphylinoid larvae it is the prothorax whichbears the most complete setal pattern. The notation and designation of setae and rows also follow theAshe-Watrous system. Abbreviations are those commonly used in their study. However, we made noattempt to homologize setae between Hydraenidae and Staphylinidae. Thus, the same setal code doesnot necessarily mean a direct homology.

RESULTS

Description of the Larva of Limnebius cordobanus d'Orchymont, 1938(Figs. 1- 24)

General body form (Figure 1) elongate, cylindrical in cross-section and nar-rowed posteriorly. Color white, head capsule, terga and urogomphi slightly yel-lowish-brown.

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40 JUAN A. DELGADO AND AGUSTfN G. SOLER

Figs. 2-4.Limnebius cordobanus, larval instar I. Head. 2: dorsal view. 3: lateral view. 4: ventralview. Abbreviations". CG: cephalic gland; Cl, clypeal setae; EC: epicranial campaniformsensilla; Ed: epicranial dorsal setae; El: epicranial lateral setae; Em: epicranial marginalsetae; FC: frontal campaniform sensillum; Fd: frontal discal setae; Fl: frontal lateral setae;Fm: frontal marginal seta; L: lateral setae; LC: lateral campaniform sensillum; T: temporalsetae; V: ventral setae.Scale = 0.1mm.

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LARVA OF LIMNEBIUS (COL.: HYDRAENIDAE) 41

Instar ITotal body length about 2.3 mm.

Head: Head capsule width: 0.38 ± 0.02 mm (Mean ± SD\ n=6). Ecdysialsutures distinct and complete from antennal fossae anteriorly to base of headposteriorly. Five stemmata are present, ocular area subdivided into 2 obliquerows. Chaetotaxy as follows and in Figures 2 to 4. Frontal region with 1 campan-iform sensillum and 5 setae arranged in 3 rows: 2 frontal dorsal setae, 2 frontallateral setae and 1 frontal marginal seta. Epicranial region with 2 campaniformsensilla, 1 epicranial gland and 10 setae arranged in 4 rows: 4 minute posteriorsetae, 2 epicranial dorsal setae, 2 epicranial lateral setae and 2 epicranial margin-al setae. Temporal region with 4 setae: Tl to T4. Lateral region with 1 campani-form sensillum and 2 setae: LI and L2. Ventral region with 2 short setae: VI andV2. No egg bursters are associated with the head capsule, although the marginaledge of the frontal region is strongly sclerotized and may be used as cutting edge.Labrum (Fig. 5) roughly semicircular, with 2 campaniform sensilla and 7 setaearranged in 2 rows: 2 labral dorsal setae and 5 labral marginal setae. Lml, Lm3and Lm5 arise dorsolaterally, Lm2 and Lm4 arise ventrolaterally. Lm4 is a verylong seta and Lm2 is bifid. Antenna (Fig. 6) with article I short, with 4 campan-iform sensilla around the apex, without setae; article II about 2.5x as long asarticle I; article III about 0.7x as long as II. Article II with 3 long setae and 3solenidia, 2 of which may be considered as sensory appendages. Article III with4 setae and 3 apical solenidia; IIS3 long. Mandibles (Fig. 7) right and left nearlyidentical in size and shape; with 2 campaniform sensilla and 2 setae. Ml long andM2 of medium length. Prostheca wide and molar area well developed. Maxilla asin Figure 8. Cardo triangular, with 1 seta. Stipes not distinctly separated frommala, surface with 4 setae and 1 campaniform sensillum. Galea and lacinia fim-briate. Palpifer consisting of crescentic sclerite at base of maxillary palpus; with1 long seta. Maxillary palpus with 3 articles, I and HI nearly equal in length,article II about 0.7x as long as I. Apical article with conspicuous, basal, digiti-form, sensory appendage on external surface; article I with 2 campaniform sen-silla, article II with 2 setae. Labium as in Figure 9, consisting of 3 sclerites;ligula short and broad, bearing papillae; palps with 2 articles, article I with 1campaniform sensillum on mesal margin. Submentum with 1 pair of setae; men-turn with 2 pairs of setae and 1 pair of campaniform sensilla. Prementum with 2pairs of setae.

Thorax: Pronotum transverse. Chaetotaxy as follows and in Figure 10: row Awith 4 setae: Al to A4; row L with 3 setae: LI to L3; row P with 4 setae: PI toP4; row Da with 1 seta: Dal; row Db with 1 seta: Dbl; row Dc with 1 seta: Del.4 campaniform sensilla are present: Cl, C2, C3 and C5; C4 absent. Mesonotumas in Figure 11; row A with 4 minute setae; row L with 3 setae: LI to L3; row Pwith 4 setae: PI to P4; rows Da, Db and Dc each with 1 seta: Dal, Dbl and Delrespectively. Campaniform sensilla Cl and C2 absent and replaced by pretergal

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42 JUAN A. DELGADO AND AGUSTJN G. SOLER

Figs. 5-9. Limnebius cordobanus, larval instar I. 5: labrum, dorsal view; 6: antenna, ventral view;7: mandible, dorsal view; 8: maxilla, ventral view; 9: labium, ventral view. Abbreviations:C: campaniform sensilla; Ld: labral dorsal setae; Cdo: cardal setae; La: seta of the lacinia;Lg: ligula; Lm: labral marginal setae; M: mandibular setae; Mnt: mentum; Pf: seta of thepalpifer; Pm: palpal setae; Pmnt: prementum; SD: sensorial digitiform appendage; Smnt:submentum; Stp: setae of the stipes; IIS (1-3), antennal solenidia of article II; HIS (1-3),antennal solenidia of article III.Scale = 0.05 mm.

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LARVA OF LIMNEBIUS (COL.: HYDRAENIDAE) 43

glands. Metanotum similar to mesonotum. Prosternum as in Figure 12; prester-num divided in 2 lateral regions, each with 2 setae; eusternum indistinctly sclero-tized; sternum consisting of central, subtriangular sclerotized area with 1 pair ofsetae. Laterosternum fused to prehypopleuron forming a sclerotized lobe anteriorto coxa, with 3 setae. Posthypopleuron consisting of slightly sclerotized lobeposterolaterad to coxa, with associated denticles or minute spines. Mesosternum(Fig. 13) differs from prosternum as follows: eusternum with 2 pairs of setae;presternal region with only 1 seta, prehypopleuron with 2 setae. Mesothoracicspiracle present, annular, with 1 seta. Metasternum similar to mesosternum. Me-tathoracic spiracle vestigial, without seta. Legs: Prothoracic leg as in Figures 14and 15. Coxa with 1 campaniform sensillum and 14 setae: 1 D, 2 Ad, 4 Al, 1 Av,

11

Figs. 10-11. Limnebius cordobanus, larval instar I. 10: pronotum; 11: mesonotum. Abbreviations:A: anterior setae; C: campaniform sensilla; Da, Db and Dc: discal setae; L: lateral setae;P: posterior setae; Pg: pretergal gland.Scale = 0.1mm.

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44 JUAN A. DELGADO AND AGUSTIN G. SOLER

12

13

Figs. 12—13. Limnebius cordobanus, larval instar I. 12: prosternum; 13: mesosternum. Abbrevia-tions: CX: coxal cavity; Pr: presternal setae; Prehy: prehypopleural setae; Pohy: posthy-popleural setae; S: spiracular seta, Sp: spiracle; St: sternal setae.Scale = 0.1 mm.

1 Pd, 4 PI and lPv. Trochanter with 7 campaniform sensilla and 8 setae: 1 Ad, 2Al, 2 Av, 1 PI, 1 Pv and 1 V. Femur with 2 campaniform sensilla and 8 setae: 1 D,1 Ad, 1A1, 2 Av, 1 Pd, 1 PI and 1 V. Tibia with 2 campaniform sensilla and 8setae: 2 D, 1 Ad, 1 Al, 1 Av, 1 Pd, 1 PI and 1 Pv. Tarsungulus with 2 minutesetae.

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LARVA OF UMNEBIVS (COL.: HYDRAENIDAE) 45

, Tb

Cx

14

Cx

Tb

15

16

Figs. 14-16. Limnebius cordobanus, larval instar I. 14: proleg, anterior view; 15: dorsal view.Abbreviations: Ad: anterodorsal setae; Al: anterolateral setae; Av: anteroventral setae;Bs: basal setae; C: campaniform sensilla; Cx: coxa; D: dorsal setae; Fm: femur; Pd:posterodorsal setae; PI: posterolateral setae; Pv: posteroventral setae; Tb: tibia; Tr:trochanter; Ts: tarsungulus; V: ventral setae; 16: abdominal tergum I. A: anterior setae;C: campaniform sensilla; DP: dorsopleural setae; L: lateral setae; P: posterior setae; Pg:pretergal gland; Sp: spiracle.Scale = 0.1 mm.

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46 JUAN A. DELGADO AND AGUSTIN G. SOLER

Abdomen: Abdominal terga are characterized in staphylinoid larvae by fur-ther shortening and widening of the sclerites. Tergum I as in Figure 16. Seta P3absent; setae Dal, Dbl and Del absent; anterior setae A2 and A3 absent. Direct-

17

21

Figs. 17-21. Limnebius cordobanus. 17: abdominal tergum I, instar III; 18: abdominal sternum I,instar I; 19: abdominal sternum I, instar III; 20: abdominal sterna II—VIII, instar I; 21:abdominal sterna II—VIII, instar III. Abbreviations: A: anterior setae; C: campaniformsensilla; D: discal setae; D': discal subprimary seta; DP: dorsopleural setae; L: lateralsetae; P: posterior setae; Pg: pretergal glands; Ps: presternal setae; Sp: spiracle; VP:ventropeural setae.Scale = 0.1 mm.

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LARVA OF LIMNEBIUS (COL.: HYDRAENIDAE) 4 7

ly below the posterolateral region of tergum I there is a pair of lateral sclerites orpleurites. The dorsopleural sclerite is located between the tergum I and the ven-tropleural sclerite. One spiracle and 2 setae are on this sclerite. The ventropleuralsclerite is located between the sternum I and the dorsopleural sclerite. Only 1seta is on this sclerite. Sternum I as in Figure 18. Seta P3 absent. Abdominalterga II-VIII similar to tergum I. Dorsopleural and ventropleural sclerites of theabdominal segments II to VIII each with 2 setae. Abdominal sterna II-VIII simi-lar to sternum I except for addition of seta D2 (Fig. 20). Setal homologies onsegments IX and X are difficult to establish. Tergum IX as in Figure 22, with 4setae and 1 pretergal gland; no spiracles detectable. Urogomphus as in Figure 23,long, with 2 articles; basal article (URI) not fused to tergum IX, with 6 setae and4 pores or campaniform sensilla. Article II (URII) short and slender, 0.3x as longas article I. URII with a long seta: AE, arising from apex. Sternum IX as inFigure 24. Abdominal segment X as in Figures 22 and 24. Anal vesicle, pseudo-pod or pygopod bearing a pair of stout hooks, and a slightly sclerotized area with3 pairs of small setae and 1 pair of campaniform sensilla.

Ins tar IISimilar to instar HI; Total body length about 3 mm. Head capsule width:0.50 ± 0.03 (Mean ± SD; n=5). Morphologically identical to third instar (Fig. 1).

Instar IIITotal body length about 4 mm. Head capsule width: 0.67 ± 0.03 (Mean ± SD;n=6). Chaetotaxy as in Figures 17,19 and 21; with following differences to instarI: abdominal tergum I-VIII with 2 additional lateral setae (Fig. 17). We interpretthe 2 new setae as DPI and DP2 located on dorsolateral sclerite in instar I. Theloss of dorsolateral sclerites in instars II and III is clearly connected with theapparition of spiracles and setae DPI and DP2 attached to tergal region of abdo-men. Abdominal sternum I with 1 additional lateral seta (Fig. 19). We interpretthis new seta as VP1 of instar I. Abdominal sternum II-VIII (Fig. 21) with 3additional setae: 2 lateral setae: VP1 and VP2 of instar I, and 1 new discal seta:D'. This last seta has been observed in other species of Limnebius and is thusconsidered as a subprimary seta. In some examined species of Limnebius morevariability was observed. We have found several cases of secondary setae ininstars other than the first.

DISCUSSION

As mentioned in the introduction the aims of this paper were to examine themorphology and chaetotaxy of the three larval instars of Limnebius and not todiscuss the phyletic relationships between Hydraenidae and other polyphaganlineages. However, in a comparison of the distribution of the primary setae andpores in Limnebius with those of other taxa, such as Aleocharinae (Ashe, 1986)

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48 JUAN A. DELGADO AND AGUSTIN G. SOLER

22

Figs. 22-24. Limnebius cordobanus, larval instar I. Segments IX and X. 22: dorsal view; 23: urogom-phus; 24: ventral view. Abbreviations: A: anterior seta; AE: apical stylus; C: campani-form sensilla; D7: indetermined discal seta; P?: indetermined posterior setae; Pg: preter-gal gland; Ps: pleurosternal seta; U: setae of article I; URI: basal article; URII: apicalarticle. ?: uncoded setae.Scale = 0.1 mm.

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LARVA OF LIMNEBIUS (COL.: HYDRAENIDAE) 4 9

or Leiodidae (Wheeler, 1990), a common pattern of mechanoreceptors can easilyappreciated among these three groups. We think that the results of our studysuggest newly a close affinity between Hydraenidae and several Staphylinoidea(see Dybas, 1976) although, of course, there are various other interpretationswich would explain this similarity as a retention of plesiomorphic characters oreven homoplasies. In this sense, chaetotaxal analysis of additional taxa might bevery useful not only for the selection of phylogenetic features, but also for thedetermination of the polarity of character states in these phyletic (cladistic) stud-ies.

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