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INTRODUCTION
Scydmaeninae is a large subfamily of Staphylinidae,
comprising nearly 4900 species in about 90 genera and
ten tribes (Grebennikov & Newton, 2009). Larval mor-
phology of ant-like stone beetles, however, is very poorly
known. To date, various authors have published descrip-
tions and illustrations of larvae of only twelve genera and
less than 0.5% of known species. Some published figures
are highly simplified and some misleading, with morpho-
logical structures misinterpreted or taxa misidentified
[e.g., a larva of the oxyteline staphylinid Syntomium
Curtis assigned to Euconnus Thomson (Schmid, 1988a;
Newton & Franz, 1998)]. Furthermore, descriptions based
on immature stages obtained ex ovo by rearing beetles are
scarce and identifications based solely on the co-
occurrence of larvae and adults may be unreliable. All
larval stages and the pupa are described only for a single
species (De Marzo, 1984) and only for one species are the
details of the chaetotaxy illustrated and described using
modern methods (Wheeler & Pakaluk, 1983). Although
larval characters of Scydmaeninae were recently used in
an extensive phylogenetic analysis (Grebennikov & New-
ton, 2009) and may be crucial for clarifying suprageneric
relationships within this subfamily, such data is scarce,
which hampers further taxonomic study.
In 1882 Tömösvary described a family Anisosphaeridae
in Thysanura for a new genus, Anisosphaera, an odd-
looking arthropod from North-Eastern Slovakia. Silvestri
(1899) placed this family in Anisosphaeridia, a new insect
order, with possible affinities to Collembola. Bergroth
(1899) recognized in Anisosphaera a larva belonging to a
scydmaenine genus, Cephennium Müller & Kunze, and
synonymized these names; an additional discussion sup-
porting this action was published by Dudich (1927). The
larvae of Cephennium were the first to be described and
are still the best known among the Scydmaeninae, with
various morphological details illustrated for four species
(Peyerimhoff, 1899; Jeannel, 1909; Scholz, 1926;
Ionesco, 1937; Paulian, 1941; Brown & Crowson, 1980;
Schmid, 1988a, b). In addition to Cephennium, in the
tribe Cephenniini there are only illustrations of the gen-
eral habitus and a few morphological details of Cephen-
nodes Reitter (as an unidentified genus by Morimoto &
Hayashi, 1986 and as Coatesia Scott by Newton, 1991).
In Eutheiini, some details are described and illustrated for
larvae of Eutheia Stephens and Veraphis Casey (Brown
& Crowson, 1980; Newton, 1991); in Cyrtoscydmini for
Stenichnus Thomson, Scydmoraphes Reitter, Neuraphes
Thomson and Euconnus (Paulian, 1941; Franz, 1965;
Brown & Crowson, 1980; Wheeler & Pakaluk, 1983;
Schmid, 1988a; Newton, 1991); in Scydmaenini for Scyd-
maenus Latreille only (Meinert, 1888; Böving & Craig-
head, 1931; Jeannel & Paulian, 1945; Brown & Crowson,
1980; Newton, 1991; Beutel & Molenda, 1997); and in
Mastigini, Clidicini and Leptomastacini larvae of one
genus in each tribe are known, respectively Palaeostigus
Newton, Clidicus Laporte and Leptomastax Pirazzoli (De
Marzo, 1983, 1984; Vit & De Marzo, 1989; O’Keefe &
Monteith, 2000; Grebennikov & Newton, 2009). Data
from the above-listed literature were used by several
Eur. J. Entomol. 109: 587–601, 2012
http://www.eje.cz/scripts/viewabstract.php?abstract=1745
ISSN 1210-5759 (print), 1802-8829 (online)
Larval morphology of Scydmaenus tarsatus and S. hellwigii, with notes
on feeding behaviour and a review of the bibliography on the preimaginal
stages of ant-like stone beetles (Coleoptera: Staphylinidae: Scydmaeninae)
PAWEà JAàOSZYēSKI1 and ALEKSANDRA KILIAN2
1Museum of Natural History, Wrocáaw University, Sienkiewicza 21, 50-335 Wrocáaw, Poland; e-mail: scydmaenus@yahoo.com
2Department of Invertebrate Biology, Evolution and Conservation, Wrocáaw University, Przybyszewskiego 63/77, 51-148 Wrocáaw,
Poland; e-mail: a.kilian@biol.uni.wroc.pl
Key words. Coleoptera, Staphylinidae, Scydmaeninae, Scydmaenini, Scydmaenus, Palaearctic, larval morphology, behaviour,
feeding
Abstract. Morphology of mature larvae of two Central European species of Scydmaenus Latreille is described and illustrated: S. (s.
str.) tarsatus Müller & Kunze and S. (Cholerus)hellwigii (Herbst). Inaccuracies in previous descriptions of S. tarsatus are discussed
and the following combination of characters is defined as diagnostic for Scydmaenus: epicranial sutures reaching posteromedian
margins of antennal insertions; presence of a single pair of stemmata and epicranial supraantennal pits; anterior row of subtriangular
teeth on epipharynx; mandibles falciform and without mesal teeth; antennomere 3 rudimentary; antennal sensory appendage sub-
conical and asymmetrical; maxilla with galea and lacinia; labium strongly constricted between mentum and prementum; thoracic ter-
gites undivided along midline; head capsule, thoracic tergites, laterotergites and abdominal segments except sternite 1 densely
setose; thoracic sternites and abdominal sternite1 largely asetose; and lack of urogomphi. We also describe the feeding behaviour of
immature S. tarsatus and demonstrate for the first time that Scydmaeninae larvae can feed on live springtails and not armoured
mites. In the introduction we provide a summary of the literature on all hitherto known preimaginal stages of Scydmaeninae.
587
authors in identification keys, reviews and summaries
(e.g., Ghilarov, 1964; Kasule, 1966; Klausnitzer, 1978,
1997; Newton & Franz, 1998, O’Keefe, 2005). Immature
stages of Plaumanniolini, Chevrolatiini and Leptoscyd-
mini remain unknown.
Data on the biology of immature Scydmaeninae, espe-
cially their behaviour and feeding preferences, are even
scarcer. Schuster (1966a, b) observed larvae of Cephen-
nium majus Reitter and C. thoracicum Müller & Kunze
feeding and carried out prey preference experiments.
These species were found to be predacious and feed
mostly on the armored mites (Oribatida), and only rarely
on Uropodina and Gamasida; cannibalistic tendencies
among the larvae were also mentioned (Schuster, 1966b).
Schmid (1988) made similar observations on immature
Cephennium,Stenichnus and Scydmoraphes. De Marzo
(1983) reared larvae of Palaeostigus pilifer (Kraatz) and
reported that the first and second instars feed exclusively
on a secretion produced by the female abdominal gland,
which was deposited along with the eggs. In the labora-
tory mature (i.e. third instar) larvae were fed on the vis-
cera of a caterpillar of Galleria sp. De Marzo is so far the
only author to provide photographic documentation of
scydmaenine larvae feeding.
Recently JaáoszyĔski (2012a) reported that adults of
Scydmaenus tarsatus Müller & Kunze and Scydmaenus
hellwigii (Herbst) can be maintained in long-term labora-
tory cultures by feeding them on soft-bodied arthropods.
Neither of these species showed any interest in oribatid or
uropodine mites, which have heavily sclerotized cuticles,
whereas both readily fed on live hypogastrurid springatils
(genus Ceratophysella Borner) and acaridid mites (genus
Rhizoglyphus Claparède). Scydmaenus hellwigii also fed
on dead springtails and S. tarsatus on dead flies, isopods
and pseudoscorpions (JaáoszyĔski, 2012b). During these
experiments, immature scydmaenine were collected
together with numerous adults of S. tarsatus and identi-
fied as conspecific with adult beetles on the basis of pre-
vious descriptions (Meinert, 1888; Brown & Crowson,
1980) and the fact that the substrate (decaying plant
matter in a garden compost heap) was inhabited only by
this species of ant-like stone beetles. During previous
rearing of S. hellwigii adults (JaáoszyĔski, 2012a) a single
larva was obtained. In the present paper the previously
inadequately described larva of S. tarsatus is redescribed,
and the larva of S. hellwigii is described for the first time.
Moreover, observations on the previously unknown
feeding behaviour of larval S. tarsatus are reported.
MATERIAL AND METHODS
Material
Larvae together with numerous adults of Scydmaenus (s. str.)
tarsatus Müller et Kunze, 1822 were collected from a large
compost heap at Wáocáawek (Central Poland) by sifting in the
last week of August 2010. No other species of ant-like stone
beetles were found in this substrate. Some larvae were immedi-
ately preserved in 75% ethanol for morphological study, six
others were placed alive in a 150 ml plastic container half-filled
with compost and transferred to the laboratory for further obser-
vations. Numerous adults of Scydmaenus (Cholerus) hellwigii
(Herbst, 1792) were collected from rotten wood and from under
loose bark of a lime tree at Wrocáaw-Wojnów (SW Poland) in
the middle of April 2010, and transported to the laboratory in a
similar manner. Scydmaenus hellwigii was reared as described
previously (JaáoszyĔski, 2012a): 20 females and 10 males were
placed in a 30 cm high, 1 l cylindrical container 3/4 filled with
moist rotten lime wood inhabited by a colony of small Isoto-
midae springtails of the genus Desoria Agassiz & Nicolet. The
container was closed except for a 50 × 2 mm ventilation
opening at the top and the substrate was dampened every two
weeks. Every two months the entire substrate was sifted and
searched for larvae and then put back into the same container
together with all the inhabitants; this culture was terminated
after six months, when a larva was found. The larva was pre-
served in 75% ethanol. Larvae of both of the species studied
were identified as last instars based on their body length,
exceeding that of adults by 20–30%.
Preparations
One larva of each species was macerated in warm 10% NaOH
after separating the head; when soft tissues had dissolved speci-
mens were washed in distilled water, slightly stained with chlo-
razol black and mounted in glycerol-gelatin. Drawings and
measurements were made at magnifications up to 600× using a
phase contrast compound microscope.
Scanning electron microscopy (SEM)
A larva of S. tarsatus was transferred from ethanol to distilled
water through a series of 70/40/20% ethanol, cleaned for 1 min
in 10% NaOH at ambient temperature, washed in water, trans-
ferred to absolute ethanol through a series of 20/40/75%
ethanol, kept for 1 h in acetone and subsequently for 1 h in hex-
amethyldisilazane. The dehydrated specimen was mounted on a
SEM stub with a carbon tab and sputter-coated with gold (Pirani
501, Edwards) and then examined using a LEO 435 VP scan-
ning electron microscope.
Light photography and image processing
Habitus images were taken using an Olympus C-750UZ
digital camera with a Raynox MSN-202 close-up lens. Image
stacks were processed using Combine ZP (Hadley, 2010). Final
image adjustments and annotations were made in Corel Photo-
Paint.
Terminology and measurements
Terminology used for the chaetotaxy follows systems pro-
posed by Wheeler & Pakaluk (1983), Wheeler (1990) and Kilian
(2007). The total body length is the sum of the lengths of the
head, thoracic and abdominal segments measured separately.
The following abbreviations are used (abdominal segments,
antennomeres and palpomeres are counted from base of abdo-
men, antenna and palp, respectively; i.e. An3L is the length of
antennomere 3): AnL – length of antennomere; AbL – length of
abdominal segment measured along midline, excluding lateral
lobes projecting caudad; measurement of segment 10 does not
include extruded anal membrane; AbW – width of abdominal
segment; ASL – length of antennal sensory appendage; HL –
length of head excluding the teeth on nasale; HW – width of
head; LPL – length of labial palpomere; MPL – length of max-
illary palpomere; MsL – mesothoracic length; MsW – mesotho-
racic width; MtL – metathoracic length; MtW – metathoracic
width; PL – prothoracic length; PW – prothoracic width.
Observations on feeding behaviour
Larvae of S. tarsatus were maintained under laboratory condi-
tions as described previously for adults (JaáoszyĔski, 2012a),
with minor modifications. Petri dishes (diameter 3 cm) half-
588
filled with plaster of Paris were used as arenas; they were pre-
conditioned by filling with moist compost for 2–3 days, then the
compost was removed and a fresh thin, pressed layer of compost
was placed on the plaster. Six larvae were observed, three in
each arena; the substrate was moistened every second day with
0.2 ml of distilled water and the Petri dishes were kept at
ambient temperature (22–24°C) and in the dark. Larvae were
provided with a mixture of various live mites belonging to
Oribatida (mostly Phthiracaridae, Galumnidae and Cara-
bodidae), Mesostigmata (Uropodina, Parasitina) and various
springtails (mostly Hypogastruridae and Tomoceridae), all col-
lected from the compost inhabited by S. tarsatus. Prey that were
not eaten were replaced every 7 days. The larvae were kept for 4
weeks and observations were made every day for long enough
to witness the entire feeding process from an attack to aban-
doning the remnants of the prey, which takes typically 1–2 h.
RESULTS
Redescription of mature larva of Scydmaenus tarsatus
Body (Figs 1–3). Total body length 2.690 mm. Elon-
gate, strongly flattened, distinctly more convex dorsally
than ventrally, nearly parallel-sided up to abdominal seg-
ment 5, abdominal segments 6–10 gradually narrowing;
pigmentation yellowish-brown; integument weakly scle-
rotized, densely setose; all setae simple (i.e., non-
modified, slender and pointed at apex); most setae on
head, thoracic and abdominal segments inserted on dis-
tinct protuberances (papillae). Tergal and sternal plates
distinct and undivided along midline. Microsculpture
absent except for fine isodiametric granulation on bases
of abdominal segments 9 and 10. Head capsule (except
for anterior part of frontoclypeal region), tergal and
sternal plates of thorax and abdomen densely covered
with fine microtrichia (Figs 5, 8).
Head (Figs 4, 5, 10–14). Prognathous, nearly as wide
as long, nearly half as wide as prothorax, broadest near
middle, distinctly flattened; HL 0.344 mm, HW 0.364
mm, HL/HW 0.94. Small, single stemma located dorso-
laterally to each antennal insertion; stemmata distinctly
darker than surrounding cuticle. Epicranial stem long,
almost half as long as cranium. Epicranial sutures nearly
straight, V-shaped, anteriorly nearly reaching antennal
insertions; accompanied by a small epicranial supra-
antennal pit. Each epicranial plate with about 30 mostly
asymmetrically distributed setae, and a dorsolateral
cluster of five dome-shaped structures (Fig. 5); frons with
about 30 mostly asymmetrically distributed setae. Ante-
rior part of frontoclypeal region (nasale) (Fig. 11) with 8
teeth of various sizes located below distinct transverse
589
Figs 1–3. Scydmaenus tarsatus, general habitus of mature larva in dorsal (1), ventral (2) and lateral (3) views. Scale bar: 0.5 mm.
marginal ridge but well-visible in dorsal view, two pairs
of fine marginal setae, three pairs of long anterior setae
and two pairs of long sub-anterior setae. Ventral side
(Fig. 12): posterior tentorial pits well-visible at base of
mouthparts; each lateromedian part of cranium (gena)
with 9–14 asymmetrically distributed setae and two pores
lateral to mouthparts, posterior part of cranium asetose.
Tentorium (Figs 13, 14) with broad and long anterior and
posterior tentorial arms and very slender and curved
dorsal tentorial arms. Tentorial bridge was not found in
our preparations.
Antenna (Fig. 15). An1L 0.034 mm, An2L 0.137 mm,
An3L 0.012 mm and ASL 0.032 mm. Antennomere 1
short, with one dorsal seta. Antennomere 2 almost 4 times
as long as 1, with long dorsal lateral seta, two long dorsal
sub-apical setae, two long apical setae, one short ventral
sub-apical seta, wide and short antero-dorsal conical sen-
sory appendage and a very small conical sensillum at its
base. Antennomere 3 very small and short with one short
basal seta and three apical pointed processes.
Labrum. Fused with frontoclypeal region. Epipharynx
(Fig. 16) with dense microtrichia directed antero-mesally.
590
Figs 4–9. Scydmaenus tarsatus. 4 – dorsal view of head, and apical part of prothorax (framed area enlarged in Fig. 5); 5 – details
of epicranial plate (magnified framed area from Fig. 4) with dome-shaped structures, microtrichia and setae; 6 – left mandible in
dorsal view, framed area magnified; 7 – pretarsus and spines of tibiotarsus; 8 – spiracle on abdominal laterotergite 4; 9 – anal mem-
brane with internal hooks. Abbreviations: ah – hook of anal membrane; ds – dome-shaped structures; m – microtrichium; s – seta; sp
– apical spine; spr – spiracle.
Mandibles (Figs 4, 6, 17). Slightly darker than body,
slightly asymmetrical, each falciform, with broad base
and slender distal part; apices blunt, mesal margin sharp,
external margin with single sub-basal seta, dorsal surface
with single sub-basal dome-shaped structure and a patch
of irregular microsculpture near base (Fig. 6).
Maxilla (Fig. 18). Cardines mesally fused to submen-
tum. Stipes broad and short, with two setae, bearing large
stipital projection with galea and lacinia. Lacinia with
dense and long apical trichia, its basal part and distal part
of galea with dense and short trichia, base of galea with
two short marginal setae; outer margin of stipital projec-
tion with one long marginal seta, its ventral surface with
two sub-basal setae. Maxillary palp inserted on distinct
palpifer with one seta; three-segmented, MP1L 0.037
mm, MP2L 0.050 mm and MP3L 0.081 mm; palpomere 1
slightly longer than wide, asetose; 2 elongate and slightly
narrowing towards apex, with two sub-median setae; 3
strongly elongate and slender, nearly straight, with single
basal seta.
Labium (Figs 19, 20). Elongate, lightly sclerotized.
Submentum fused to cardines, with one basal pair of
setae. Mentum narrowing anteriorly, with two sub-
median pairs of setae. Prementum broadening anteriorly,
indistinctly demarcated from mentum, with broadly emar-
ginated anterior margin, with one lateral pair of sub-
anterior ventral setae and two lateral pairs of anterior
dorsal setae; ligula absent. Labial palps two-segmented,
LP1L 0.033 mm and LP2L 0.028 mm; palpomere 1 sub-
cylindrical and asetose, 2 narrower than 1 but similar in
length, sub-conical. Hypopharynx (Fig. 20) with two
transverse rows of fine microtrichia.
Legs (Figs 21–23). Five-segmented, slender, long,
without microsculpture. Coxa elongate, with 11–13 setae
of various lengths. Trochanter strongly elongate, on each
leg with four moderately long setae distributed sub-
591
Figs 10–14. Scydmaenus tarsatus. 10 – head in dorsal view; 11 – nasale in dorsal view; 12 – head with mouthparts in ventral view;
13 – tentorium in dorsal view; 14 – tentorium in lateral view. Abbreviations: a1–3 – anterior setae; ata – anterior tentorial arm; cd –
cardo; dta – dorsal tentorial arm; ep – epicranial pit; es – epicranial suture, est – epicranial stem; lp – labial palp; m1–2 – marginal
setae; md – mandible; me – mentum; mp – maxillary palp; p – pore; pm – prementum; pta – posterior tentorial arm; ptp – posterior
tentorial pits; s1, s2 – sub-anterior setae; sm – submentum; st – stemma; stp – stipes. Scale bars: 10, 12–14: 0.1 mm; 11: 0.05 mm.
medially and apically and one strikingly long subapical
seta. Femur strongly elongate, on each leg with 5–6 setae.
Tibiotarsus strongly elongate, as long as femur or slightly
longer (metathoracic legs), on each leg with 11 setae and
fine apical spines (Fig. 7). Pretarsus (Fig. 7) long and
curved, pointed, with two setae.
Prothorax (Figs 24, 27). Transverse, expanded later-
ally; PL 0.344 mm, PW 0.657 mm and PL/PW 0.523.
Pronotum (Fig. 24) with convex tergal plate occupying
posterior half of dorsum and large, convex anterior part;
setae distributed asymmetrically, partly forming irregular
transverse rows, setae subequal in length and diameter
except 2–3 slightly thicker and longer setae located on
each lateroposterior corner of pronotum; ca. 80 setae on
anterior part, ca. 110 on tergal plate; laterotergite with ca.
50 setae along lateral margin, ca. 12 dorsal setae and ca.
20 ventral setae of various sizes. Basisternum (Fig. 27)
with five setae forming arcuate transverse row. Pleurites
(Fig. 27) with one seta on episternum and one seta on epi-
meron.
Mesothorax (Figs 25, 28). Transverse, expanded later-
ally; MsL 0.294 mm, MsW 0.734 mm and MsL/MsW
0.400. Mesonotum (Fig. 25) with tergal plate occupying
3/4 of its posterior part; setae distributed asymmetrically,
partly forming irregular transverse rows, setae subequal
in length and diameter except two slightly thicker and
longer setae located on each lateroposterior corner and
two pairs of short and thin anterolateral pretergal setae;
tergal plate with ca. 90 setae; each laterotergite with ca.
12 dorsal setae, ca. 15 ventral and ca. 30–40 lateral setae.
Median part of sternum (Fig. 28) with three pairs of setae.
Episternum with two setae; epimeron asetose.
Metathorax (Figs 26, 29). Transverse, expanded later-
ally; MtL 0.269 mm, MtW 0.794 and MtL/MtW 0.339.
Metanotum (Fig. 26) with tergal plate occupying 2/3 of
its posterior part, narrower than mesonotal tergal plate.
Setae similar to those on mesonotum (ca. 80 setae on
tergal plate); median part of sternite (Fig. 29) with
irregular transverse row of 7 setae; episternum with two
setae; epimeron with one seta.
592
Figs 15–20. Scydmaenus tarsatus. 15 – right antenna in dorsal view; 16 – epipharynx; 17 – mandibles in dorsal view; 18 – left
maxilla in ventral view; 19 – labium in ventral view; 20 – hypopharynx. Abbreviations: an1–3 – antennomeres 1–3; cs – conical sen-
sillum; ga – galea; la – lacinia; hph – hypopharynx; lp1–2 – labial palpomere 1–2; me – mentum; mp1–3 – maxillary palpomere 1–3;
pf – palpifer; pm – prementum; SA – sensory appendage; stp – stipes; stpp – stipital projection. Scale bars: 0.05 mm.
Figs 21–23. Scydmaenus tarsatus. 21 – left prothoracic leg in
posterior view; 22 – right mesothoracic leg in posterior view; 23
– right metathoracic leg in posterior view. Abbreviations: cx –
coxa; fe – femur; pr – pretarsus; sp – spine; ti – tibiotarsus; tr –
trochanter. Scale bars: 0.05 mm.
Abdominal tergites 1–8 (Figs 30–34, 40–42). Tergites
strongly transverse, expanded laterally, tergal plates dis-
tinct only on tergites 1–4 (Figs 30–33); tergites 6–8 (Figs
40–42) gradually narrowing and becoming longer. Ab1L
0.139, Ab1W 0.750 mm; Ab1L/Ab1W 0.185; Ab2L
0.151 mm, Ab2W 0.800 mm, Ab2L/Ab2W 0.189; Ab3L
0.150 mm, Ab3W 0.812 mm, Ab3L/Ab3W 0.185; Ab4L
0.133, Ab4W 0.781, Ab4L/Ab4W 0.170; Ab5L 0.147
mm, Ab5W 0.762, Ab5L/Ab5W 0.193; Ab6L 0.128 mm,
Ab6W 0.687, Ab6L/Ab6W 0.186; Ab7L 0.145 mm,
Ab7W 0.594, Ab7L/Ab7W 0.244; Ab8L 0.145 mm,
Ab8W 0.456 mm and Ab8L/Ab8W 0.318. Setae
numerous and forming three irregular, transverse rows,
subequal in length and diameter, with two exceptions:
two slightly thicker and longer setae located on each
lateroposterior corner of laterotergite, and 1–2 pairs of
short and thin anterolateral pretergal setae. Tergites 1–4
each with ca. 50 setae on tergal plate; each laterotergite
with ca. five dorsal, ca. 15 lateral and 4–10 ventral setae;
pretergal setae: two pairs on tergite 1, one pair on tergites
2, 4–6, absent on tergites 3, 7–8. Tergites 5–8 with
decreasing number of setae (40–28), each laterotergite
with five dorsal, ca. 10 lateral and five ventral setae.
Abdominal sternites 1–8 (Figs 35–39, 43–45).
Strongly transverse, sternal plates distinct. Sternite 1 (Fig.
35) with scarce setae: one pair of setae in middle and two
pairs postero-laterally. Sternites 2–8 (Figs 36–39, 43–45)
with numerous setae in irregular, transverse rows: ante-
rior row with 6–12 setae, posterior row with 8–10 setae,
except sternite 8 with 36 setae forming distinct lateral
groups.
Abdominal segment 9 (Figs 46, 47). Subtrapezoidal,
slightly wider than long, narrowing caudad and with
rounded posterior margin; Ab9L 0.200 mm, Ab9W 0.272
mm and Ab9L/Ab9W 0.735; tergal plate (Fig. 46) indis-
tinct; setae numerous (ca. 50) and distributed along lateral
and posterior margin; median part largely asetose. Ster-
nite (Fig. 47) similar in shape, with ca. 60 setae irregu-
larly distributed on median and lateral parts, apical part
only with marginal setae.
Abdominal segment 10 (Figs 9, 48, 49). Subconical,
distinctly longer than wide and slightly narrowing posteri-
orly; Ab10L 0.098 mm, Ab10W 0.90 mm and
Ab10L/Ab10W 1.089; tergite (Fig. 48) with 8–9 long
593
Figs 46–49. Scydmaenus tarsatus, abdominal segments 9–10
in dorsal (46, 48) and ventral (47, 49) views. 46–47 – segment
9; 48–49 – segment 10. Scale bar: 46–47: 0.2 mm; 48–49: 0.1
mm.
Figs 30–39. Scydmaenus tarsatus, abdominal segments 1–5 in
dorsal (30–34) and ventral (35–39) views. 30, 35 – segment 1;
31, 36 – segment 2; 32, 37 – segment 3; 33, 38 – segment 4; 34,
39 – segment 5. Scale bar: 0.2 mm.
Figs 24–29. Scydmaenus tarsatus, thoracic segments in dorsal
(24–26) and ventral (27 –29) views. 24, 27 – prothorax; 25, 28 –
mesothorax; 26, 29 – metathorax. Scale bar: 0.2 mm.
Figs 40–45. Scydmaenus tarsatus, abdominal segments 6–8 in
dorsal (40–42) and ventral (43–45) views. 40, 43 – segment 6;
41, 44 – segment 7; 42, 45 – segment 8. Scale bar: 0.2 mm.
setae on each side, with largely asetose posterior and
median part; sternite (Fig. 49) with 12 long setae on each
side, with particularly long posterior and median setae.
Anal membrane with ca. 14 ventral, darkly sclerotized
hooks inside (Fig. 9) and five ventral marginal pores.
Spiracles (Figs 8, 28, 35–39, 43–45). Annular, nine
pairs of ventrolateral spiracles: one on mesothorax, on
anterior protuberance of laterotergite, with one seta at
posterior margin of spiracular stalk, eight pairs of spira-
cles on abdominal segments 1–8 located sub-medially or
posteriorly on laterotergites.
Feeding behaviour of immature Scydmaenus tarsatus
The only prey observed to be attacked and eaten by
larvae of S. tarsatus were live springtails: Ceratophysella
denticulata (Bagnall, 1941) (Hypogastruridae) (Figs
50–53) and Tomocerus sp. (Tomoceridae) (Figs 54–55).
Although after 7 days some mites were found dead, their
bodies were apparently intact and undamaged. Larvae did
not show any interest in mites present in the arena, but
readily attacked springtails. All Collembola attacked and
devoured were smaller than the Scydmaenus larvae, typi-
cally about 1–1.5 mm in length. Beetle larvae actively
searched for prey by patrolling the arena and the spring-
tails they encountered were usually attacked from behind
or from the side. During four weeks, the entire feeding
from attack to abandoning the remains of the prey was
observed 22 times. In each case the first contact with
maxillary palps was immediately followed by a quick grip
with mandibles; the feeding time depended on the size of
prey captured and was from about 20 to 50 min. During
this time the prey showed irregular movements for several
minutes. The Scydmaenus larvae frequently changed their
grip on their prey during feeding (often turning the
594
Figs 50–55. Scydmaenus tarsatus feeding on Ceratophysella (50–53) and Tomocerus (54–55) springtails.
springtail upside down and consuming it from the ventral
surface) and chewing intensively with their mandibles.
The prey was devoured nearly completely; the abandoned
remains were not recognizable as a springtail.
Description of mature larva of Scydmaenus hellwigii
Body (Fig. 56). Total body length 2.31 mm. Strongly
elongate, sub-cylindrical and only slightly flattened, more
convex dorsally than ventrally, nearly parallel-sided up to
abdominal segment 2, remaining part of abdomen gradu-
ally narrowing; pigmentation creamy-white; integument
weakly sclerotized, densely setose, all setae simple (i.e.,
non-modified, slender and pointed at apex) and with
simple (i.e., not papillate) insertions. Tergal and sternal
plates indistinct and undivided along midline. Micro-
sculpture absent except for fine isodiametric granulation
on bases of abdominal segments 9 and 10 and most of the
surface of abdominal sternite 9. There are no microtrichia
on the body segments.
595
Fig. 56. Scydmaenus hellwigii, habitus of larva in dorsal
view. Scale bar: 0.5 mm.
Figs 57–59. Scydmaenus hellwigii. 57 – head in dorsal view;
58 – nasale in dorsal view; 59 –head with mouthparts in ventral
view. Abbreviations as in Figs 10–14. Scale bars: 57, 59:
0.1 mm; 58: 0.05 mm.
Head (Figs 57–59). Prognathous, nearly as wide as
long, only slightly narrower than prothorax, broadest
slightly posterior to middle, distinctly flattened; HL 0.264
mm, HW 0.269 mm and HL/HW 0.981. Small, single
stemma located dorso-laterally and posteriorly to each
antennal insertion; stemmata distinctly darker than sur-
rounding cuticle. Epicranial stem long, about half as long
as cranium. Epicranial sutures nearly straight, v-shaped,
anteriorly nearly reaching antennal insertion, accompa-
nied by a small epicranial supra-antennal pit. Each epicra-
nial plate with 40–50 mostly asymmetrically distributed
setae; frons with ca. 25 setae. Anterior part of fronto-
clypeal region (nasale) (Fig. 58) with five large teeth
located below distinct transverse marginal ridge and not
visible in dorsal view, one pair of fine marginal setae,
three pairs of long anterior setae and two pairs of long
sub-anterior setae. Ventral side (Fig. 59): tentorial pits not
visible; each latero-median part of cranium (gena) with
ca. 10 asymmetrically distributed setae and one pore lat-
eral to mouthparts; posterior part of cranium asetose. Ten-
torium similar to that in S. tarsatus (not shown).
Antenna (Fig. 60). Three-segmented, An1L 0.032 mm,
An2L 0.087 mm, An3L 0.006 mm and ASL 0.018 mm.
Antennomere 1 short, with one seta. Antennomere 2
almost 3 times as long as 1, with sub-median dorsal pore,
long dorsal medio-lateral seta, two long dorsal subapical
setae, two long apical setae, one short ventral sub-apical
seta, wide and short antero-dorsal conical sensory
appendage and a very small conical sensillum at its base.
Antennomere 3 very short with long sub-basal seta and
two apical pointed processes.
Labrum. Fused with frontoclypeal region. Epipharynx
(Fig. 61) with dense antero-mesally directed microtrichia.
Mandibles (Fig. 62). Slightly darker than body, sym-
metrical, each falciform, with broad base and slender
596
Figs 60–64. Scydmaenus hellwigii. 60 – right antenna in dorsal view; 61 – epipharynx; 62 – mandibles in dorsal view; 63 – right
maxilla in ventral view; 64 – labium in ventral view. Abbreviations as in Figs 15–20. Scale bars: 0.05 mm.
Figs 65–67. Scydmaenus hellwigii. 65 – right prothoracic leg
in posterior view; 66 – left mesothoracic leg in posterior view;
67 – left metathoracic leg in posterior view. Abbreviations as in
Figs 21–23. Scale bars: 0.05 mm.
distal part; apices pointed, mesal margin sharp, external
margin with single sub-basal seta, dorsal surface with
single sub-basal dome-shaped structure.
Maxilla (Fig. 63). Cardines mesally fused to submen-
tum, each with one seta. Stipes with two setae and large
projection bearing lacinia and galea. Lacinia with dense,
long apical trichia, its basal part and nearly entire surface
of galea with sparser and shorter trichia; base of galea
with two short ventral setae; outer margin of stipital pro-
jection with one marginal seta, ventral surface of stipital
projection with one sub-basal seta. Maxillary palp three-
segmented, inserted on distinct palpifer with one seta;
MP1L 0.031 mm, MP2L 0.051 mm and MP3L 0.055 mm;
palpomere 1 short and nearly as long as broad, asetose; 2
strongly elongate and narrowing towards apex, with two
sub-median setae; 3 strongly elongate, slender, slightly
curved, asetose.
Labium (Fig. 64). Elongate, lightly sclerotized. Sub-
mentum fused with cardines, with one basal pair of setae.
Mentum slightly narrowing anteriorly, with two sub-
median pairs of setae. Prementum broadening anteriorly,
indistinctly demarcated from mentum, with shallowly
emarginated anterior margin, with two lateral pairs of
anterior setae; ligula absent. Labial palps two-segmented,
LP1L 0.020 mm and LP2L 0.033 mm; palpomere 1 sub-
cylindrical and asetose, 2 narrower and much longer than
1, subconical. Hypopharynx (not shown) as in S. tarsatus.
Legs (Figs 65–67). Five-segmented, slender, long,
without microscupture. Coxa elongate, with 9–12 setae of
various sizes. Trochanter strongly elongate, on each leg
with 6–8 setae of various lengths distributed sub-medially
and apically and one strikingly long subapical seta. Femur
strongly elongate, on each leg with seven setae. Tibio-
tarsus strongly elongate, as long as femur or longer
(metathoracic legs), with 9–11 setae and fine apical
spines. Pretarsus long and curved, pointed, with two
setae.
Prothorax (Figs 68, 71). Transverse, not expanded lat-
erally; PL 0.221 mm, PW 0.326 mm and PL/PW 0.678.
Setae on pronotum (Fig. 68) numerous (77–86), distrib-
uted mostly asymmetrically, subequal in size except two
pairs of long lateral setae. Basisternum (Fig. 71) with one
pair of setae. Pleurites (Fig. 71) with two setae on each
episternum, one seta on epimeron and one sub-median
seta on laterotergite.
Mesothorax (Figs 69, 72). Transverse, not expanded
laterally; MsL 0.169 mm, MsW 0.358 mm and
MsL/MsW 0.472; setae on mesonotum (Fig. 69)
numerous (ca. 44) and arranged in several irregular trans-
verse rows, most setae subequal in size except for 2–3
pairs of minute antero-lateral setae and enlarged lateral
(marginal) setae, of which one pair is particularly long
and thick. Anterior sternal part (Fig. 72) with two pairs of
median setae. Each epimeron (Fig. 72) with one seta.
Metathorax (Figs 70, 73). Transverse, not expanded
laterally; MtL 0.132 mm, MtW 0.347 mm and MtL/MtW
0.380. Setae on metanotum (Fig. 70) and sternum (Fig.
73) similar to those on mesothorax.
Abdominal tergites 1–8 (Figs 74–78, 84–86). Tergites
transverse (1–5 strongly, 6–7 distinctly, 8 slightly), not
expanded laterally, tergal plates indistinct; tergites 1–5
(Figs 74–78) subequal in width, tergites 6–8 (Figs 84–86)
gradually narrowing and becoming longer. Ab1L 0.079
mm, Ab1W 0.379 mm, Ab1L/Ab1W 0.208; Ab2L 0.096
mm, Ab2W 0.389 mm, Ab2L/Ab2W 0.247; Ab3L 0.097
mm, Ab3W 0.379 mm, Ab3L/Ab3W 0.256; Ab4L 0.081
mm, Ab4W 0.379 mm, Ab4L/Ab4W 0.214; Ab5L 0.129
mm, Ab5W 0.379 mm, Ab5L/Ab5W 0.340; Ab6L 0.213
mm, Ab6W 0.337, Ab6L/Ab6W 0.633; Ab7L 0.218 mm,
Ab7W 0.305 mm, Ab7L/Ab7W 0.715; Ab8L 0.205,
Ab8W 0.242 and Ab8L/Ab8W 0.849. Setae numerous, on
each tergite arranged in three irregular, transverse, poste-
rior rows; lateral (marginal) setae distinctly longer and
thicker than dorsal ones. Tergites 1–5 with ca. 50 setae,
597
Figs 68–73. Scydmaenus hellwigii, thoracic segments in
dorsal (68–70) and ventral (71–73) views. 68, 71 – prothorax;
69, 72 – mesothorax; 70, 73 – metathorax. Scale bar: 0.2 mm.
Figs 74–83. Scydmaenus hellwigii, abdominal segments 1–5
in dorsal (74–78) and ventral (79–83) views. 74, 79 – segment
1; 75, 80 – segment 2; 76, 81 – segment 3; 77, 82 – segment 4;
78, 83 – segment 5. Scale bar: 0.2 mm.
tergites 6–8 with ca. 40 setae; laterotergites with 3–4 lat-
eral (marginal) setae and 2–5 ventral setae.
Abdominal sternites 1–8 (Figs 79–83, 87–89). Trans-
verse, sternal plates distinct. Sternite 1 (Fig. 79) with few
setae: two pairs of setae in middle and 2–5 setae postero-
laterally on each side. Sternites 2–8 (Figs 80–83, 87–89)
with three irregular transverse rows of setae: anterior row
with 7–10 short setae, median row of 4–8 long setae and
posterior row of 9–18 setae of various lengths. Postero-
lateral and postero-submedian pairs of setae on sternites
5–8 distinctly longer than remaining setae.
Abdominal segment 9 (Figs 90, 91). Suboval with dis-
tinct sub-basal constriction, distinctly elongate; Ab9L
0.206 mm, Ab9W 0.131 mm and Ab9L/Ab9W 1.572;
tergal plate (Fig. 90) indistinct; setae numerous (52) and
subequal in size, distributed in seven irregular rows on
posterior 2/3 of tergite. Sternite (Fig. 91) with 36 setae of
various lengths distributed asymmetrically in median
parts of sternite.
Abdominal segment 10 (Figs 92, 93). Sub-conical,
strongly elongate, slightly constricted in sub-basal region
and strongly narrowing posteriorly; Ab10L 0.200 mm,
Ab10W 0.075 mm and Ab10L/Ab10W 2.667. Setae on
tergite (Fig. 92) numerous (26) and subequal in length,
occupying posterior 3/4 of tergite; sternite (Fig. 93) with
25 setae asymmetrically distributed laterally and in three
irregular transverse rows, basal and apical parts of sternite
asetose. There are no hooks on the anal membrane.
Spiracles (Figs 71, 75, 77, 79, 81, 83, 85, 87, 89).
Annular, nine pairs of ventro-lateral spiracles: one on
mesothorax, on prominent anterior protuberance of
laterotergite, with group of fine trichia and one mesal
seta; and eight pairs of spiracles postero-laterally on
abdominal segments 1–8.
DISCUSSION
The larva of Scydmaenus tarsatus was described by
Meinert (1888) and his illustrations were redrawn several
times (e.g., Ganglbauer, 1899; Larsson, 1941; Kühnelt,
1961; Ghilarov, 1964; Klausnitzer, 1978), in some cases
losing or even gaining some details. Meinert’s description
(in Danish) is rather brief, but the genus can be deter-
mined using the figures, which are fairly accurate except
that showing the stipital projection with a single apical
lobe, instead of the two distinct lobes reported in the pre-
sent study. Some additional details provided later by
Brown & Crowson (1980) disagree with the previous
description. These authors state that the ocelli are absent,
the stipital projection is illustrated with a simple subtrian-
gular apex and the pygopod (i.e., the abdominal segment
10) with a distinct pair of lateral projections, apparently
inserted on the extruded anal membrane. The present
study confirms Meinert’s observations concerning the
presence of a pair of small stemmata (Fig. 10) and the
absence of any lateral lobes or projections on the
abdominal apex (Figs 48, 49). Moreover, the nasale illus-
trated by Brown & Crowson is devoid of teeth, while in a
slide preparation and in SEM images a row of small sub-
triangular teeth can be seen (Figs 4, 10, 11).
The larva of Scydmaenus tarsatus is unique among
known Scydmaeninae in having distinctly separated and
strongly expanded lateral parts on thoracic and abdominal
segments, the protergite with a large anterior part sepa-
rated from the tergal plate, the nasale with a row of teeth
598
Figs 84–89. Scydmaenus hellwigii, abdominal segments 6–8
in dorsal (84–86) and ventral (87–89) views. 84, 87 – segment
6; 85, 88 – segment 7; 86, 89 – segment 8. Scale bar: 0.2 mm.
Figs 90–93. Scydmaenus hellwigii, abdominal segments 9–10
in dorsal (90, 92) and ventral (91, 93) views. 90–91 – segment
9; 92–93 – segment 10. Scale bar: 0.1 mm.
visible in dorsal view, a very large number of setae that
are mostly asymmetrically distributed and inserted on dis-
tinct papillae. In general body form this larva highly
resembles immature Scydmaenus tachyoryctidis (Jeannel
& Paulian, 1944) and S. longicollis (Böving & Craighead,
1953). However, the newly described larva of Scyd-
maenus hellwigii (Fig. 56) strikingly differs from the
nearly onisciform S. tarsatus in having a more cylindrical
body, without expanded and demarcated lateral parts on
the segments, its setae have simple (i.e., not papillate)
insertions and abdominal segment 9 is strongly elongate,
not transverse (major differences are listed in Table 1).
The subcylindrical body shape resembles that of a cyr-
toscydmine species, Stenichnus turbatus Casey (Wheeler
& Pakaluk, 1983), but Scydmaenus hellwigii clearly dif-
fers in a number of structures, e.g., medially undivided
thoracic tergites (divided in Stenichnus), much denser and
asymmetrically distributed setae, anterior parts of epicra-
nial sutures adjacent to posteromedian (and not anterome-
sal) margins of antennal insertions, the antennomere 3
rudimentary (and not nearly half as long as 2) and the two
lobes of maxilla with dense microtrichia (and not a single
apical lobe bearing only several robust setae). The larva
of Scydmaenus hellwigii shares many more characters
with S. tarsatus than with any of the known immatures of
species of Eutheiini, Cephenniini, Cyrtoscydmini, Mas-
tigini, Leptomastacini or Clidicini. Larvae of S. tarsatus
and S. hellwigii can be distinguished from all remaining
Scydmaeninae by the following set of characters, diag-
nostic for Scydmaenus: the epicranial sutures reach pos-
teromedian margins of antennal insertions; the presence
of epicranial supra-antennal pits and a single pair of stem-
mata; the anterior margin of epipharynx with a row of
subtriangular teeth; the mandibles falciform, without
599
absentpresenthooks on anal membrane25.
2 pairs in middle, 2–5 pairs posterolaterally1 pair in middle, 2 pairs posterolaterallysetae on abdominal sternite 1 24.
strongly elongate, strongly narrowing
caudad
slightly elongate, weakly narrowing
caudad
abdominal tergite 1023.
distally on sternite 9ventrally on sternite 9articulation between abdominal
segments 9–10
22.
sub-oval, elongatesubtrapezoidal, transverseabdominal tergite 921.
indistinctdistinct on tergites 1–4abdominal tergal plates20.
episternum asetose; epimeron 1 setaepisternum 2 setae; mesepimeron asetose;
metepimeron 1 seta
pleural setae on meso- and
metathorax
19.
2 pairs3 pairssetae on median part of meso- and
metasternum
18.
anterior part not separatedanterior part large, convex, separated from
tergal plate
pronotum17.
only slightly wider than headnearly twice as wide as head prothorax16.
presentabsentlong lateral setae on thoracic seg-
ments
15.
less numerous, with simple insertionsmore numerous, with papillate insertionsdorsal setae on all body segments14.
lp2 longer than lp1lp2 shorter than lp1labial palp13.
broadly fused to submentumnarrowly fused to submentumcardo12.
absentpresentsubbasal mesal seta on stipital projec-
tion
11.
mp1 sub-quadrate; mp3 nearly as long as
mp2
mp1 elongate; mp3 longer than mp2maxillary palp10.
shorter, 2× An3Llonger, 3× An3Lantennal sensory appendage9.
with 2 apical processes and long sub-basal
seta
with 3 apical processes and short sub-basal
seta
antennomere 38.
stouter, 2.5× An1Lslender, 4× An1Lantennomere 27.
absentpresentdome-shaped structures on epicranial
plates
6.
indiscerniblepresentposterior tentorial pits5.
1 pair of marginal setae2 pairs of marginal setaesetae on nasale 4.
5 teeth visible only in ventral view8 teeth visible in dorsal view nasale3.
absentpresentmicrotrichia on body segments2.
sub-cylindrical, not expanded laterallynearly onisciform, flattened and expanded
laterally
body form1.
S. hellwigiiS. tarsatusCharacter
TABLE 1. Major morphological differences between larvae of S. tarsatus and S. hellwigii.
additional teeth on the mesal margin; the antennomere 3
rudimentary; the antennal sensory appendage sub-conical
and asymmetrical; the maxillae with two lobes; the
labium strongly constricted between the mentum and pre-
mentum; thoracic tergites undivided along midline; the
head, thoracic tergites, laterotergites and abdominal seg-
ments except sternite 1 densely setose; thoracic sternites
and abdominal sternite 1 largely asetose; and lack of uro-
gomphi.
Scydmaenus is a highly diverse, speciose and cosmo-
politan genus divided into about 30 subgenera (Newton &
Franz, 1998); it is even more heterogeneous than Eucon-
nus, the largest and taxonomically most challenging scyd-
maenine genus. JaáoszyĔski et al. (2012) recently demon-
strated that adults of S. tarsatus and S. hellwigii, although
externally similar, differ strikingly in the architecture of
their cephalic central nervous system; also structures on
the metaventrite are clearly different between the nomino-
typical subgenus of Scydmaenus and Cholerus (Jaáo-
szyĔski, unpubl. observ.). It is therefore not surprising
that the larvae of these species differ significantly.
This study, for the first time, demonstrates that scyd-
maenine larvae can feed on Collembola. Under laboratory
conditions, the larvae of S. tarsatus showed no interest in
mites and fed only on springtails. JaáoszyĔski (2012a, b)
reports that adults of this species ignore armoured
oribatid and uropodine mites, but feed readily on soft-
bodied acaridids and springtails (including Ceratophy-
sella and Tomocerus, accepted also by the larvae) and
scavenge various dead arthropods. Although their behav-
iour or food preferences may have been affected by labo-
ratory conditions and in nature they may prefer other
prey, it is clear that both adults and larvae of S. tarsatus
are capable of capturing and feeding on springtails. Since
the mouthparts of adult and immature Scydmaenus show
a number of differences, this observation is important for
understanding the evolution of morphological adaptations
for feeding on a particular kind of prey, characteristic of
some Scydmaeninae [e.g., Cephenniini (Schuster, 1966a,
b; Schmid, 1988a, b)].
ACKNOWLEDGEMENTS. We are indebted to J. RadwaĔski
(Department of Invertebrate Zoology, Institute of Systematics
and Evolution of Animals of Polish Acad. Sci., Kraków) for
identifying the Collembola and to R. Ruta and L. Borowiec
(Department of Biodiversity and Evolutionary Taxonomy, Uni-
versity of Wrocáaw, Wrocáaw) for their help with collecting
adults of S. hellwigii.
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Received February 20, 2012; revised and accepted April 20, 2012
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