Biology of the Stag Beetle: " de lo poco conocido y lo mucho por conocer"
(version 3.1, 2005) (See Spanish version for figures)
Working Group on Iberian Lucanidae (S.E.A.), P. O. Box 385, 33400 Avilés - Asturias (Spain)
The Stag Beetle,
Lucanus cervus (L.), is "no doubt, the most beautiful and
representative [...] beetle" (Rodríguez, 1989). Sentences
like the previous one use to open popular articles about Stag Beetle.
And afterwards, those articles usually proceed to review what is told
to be the well known biology of such a famous species.
In fact, there
are lots of things we do not know about basic aspects of the biology
of this conspicuous and emblematic beetle. Until very recently, the
few available scientific articles dealing with its life cycle were mostly
anecdotical descriptions with little or null rigorous quantification.
This is even more surprising considering that the Stag Beetle is a such
conspicuous, popular and threatened species. To be honest, panic-monger
statements such as According all the indications, its extinction
is assured within fifty years." (Huerta & Rodríguez,
1988; Rodríguez, 1989) are not based in any rigorous study.
the first version of this article was written in 1996 a considerable
quantitative and qualitative change has occurred in the amount of biological
data bout the Stag Beetle. Aside of the clasic monography by Palm (1959),
the best available summary about the biology of Middle Europe Lucanidae
is Klausnitzer (1995), which provides some general quantitative data
in support of many of the topics mentioned. Recently, a national survey
has been carried out in the United Kingdom (Percy et al., 2000), with
plentiful biological information. In addition, Sprecher-Uebersax (2001)
has finished her PhD thesis about the biology of L. cervus in
the sourroundings of Basel (Switzerland). Deborah Harvey (UK) is also
carrying out a Ph D on the Stag Beetle.
This paper summarizes the available biological data about Stag Beetle and points out to some gaps in such knowledge. Its main goal is to make obvious the need of getting a deeper knowledge of the biology of this beetle. Natural history provides a good start but is not enough; detailed quantitative research is also needed about the biology and ecology of the Stag Beetle. Only with such a knowledge its conservation can be seriously addressed.
Sexual dimorphism is remarkable in Stag Beetle and is the likely origin of its Spanish common name, Ciervo Volante (Flying Stag). Males, aside of being bigger than females, have very developed mandibles. It is considered as the biggest beetle in Europe. Plates and pictures of huge specimens included in many popular articles give us the typical image of this beetle but hide a considerable variation in the body size of the individuals. Total length ranges 30 - 90 mm in males (included mandibles) and 28 - 54 mm in females (Lacroix, 1968, 1969; Clark, 1977; Harvey & Gange, 2003 a).
Morphological variation is not limited to body size but includes some details of mandible shape and the number of lamellae on each antenna (Colas, 1962; Lacroix, 1968; Mal, 1972). This has prompted the distinction of several forms or varieties (van Roon, 1910; see Baraud, 1993 for a recent review). Of doubtful taxonomic value, several authors consider they should be abandoned (Español, 1973). López-Colón (2000), in his review of this species for the Iberian peninsula, has ignored such excesive subdivision in forms and varieties. Notwithstanding, this morphological variability is also present in other species of this family (Arrow, 1937; Otte & Stayman, 1979; Baraud, 1993) and is very interesting from other points of view, as we will see below, when talking about reproduction of Stag Beetle.
among xylophagous beetle species within a same wood piece has been reported
(Simandl, 1993). It is also known that each Lucanid species utilizes
a different portion of a tree (Szujecki, 1987). In Sweden, Palm (1959)
reported higher frequency of Stag Beetle in stumps above 20 cm diameter,
although larvae were also found in logs and dead standing trees. Within
the trunks, larvae were mainly present in the roots and in lesser degree
in the aerial part; no presence was detected in the branches (Palm,
1959). According Español (1973) larvae come quickly into the
wood and use to stay in the underground portion of the stumps. de Ligondes
(1959) found larvae burrowing within a stump. Jirí Simandl (pers.
comm.) states that larvae live free within the soil, in the contact
zone between the humus and the rotten wood. Our observations of stumps
of different tree species, larvae were found underground and although
the wood had been burrowed, larvae were also present in the ground around
the roots. Sprecher-Uebersax (2001) also documents this pattern. Sprecher
(2003) reports that young larvae feed close to the main roots of stumps,
while older larvae move further away from the roots. Murria Beltrán
et al. (2004) found larvae in the mould which accumulates in the base
of holes of Quercus cerrioides. Unfortunately, additional literature
that could shed light on this topic is written in Polish or Russian,
and out of our reach (Mamaev & Solokov, 1960; Pawlowski, 1961, quoted
in Szujecki, 1987).
Studies on the
succession of organisms involved in wood decay describe Lucanids as
appearing in the middle or late phases of this proccess, about five
years after tree death (ranging 1 - 10 years, depending on the author:
Dajoz, 1974; Szujecki, 1987). Once more, the few studies quoting Stag
Beetle are written in Russian or Polish. In general, Lucanidae are not
considered as forest pests (Palm, 1959).
Eggs hatch within
two to four weeks (Baraud, 1993; Harvey & Gange, 2003 b). Larval
life duration is variable, from one to seven years depending on the
author (Palm, 1959; Paulian, 1988; Baraud, 1993; Drake, 1994). This
slow development is due, on one hand, to the low nutritive quality of
rotten wood (low nitrogen content) and, on the other hand, to the large
size which must be achieved at maturity. Although it is assumed that
all Scarabaeoidea and Lucanoidea have three larval instars (Stehr, 1991)
and Klausnitzer (1995) gives measurements of the head capsules for those
three instars, Harvey & Gange (2003, b) have found that L. cervus
has 5-6 larval instars. Duration of each instar is unknown, although
Sprecher-Uebersax (2001) has documented the growth rate of the first
larval instars. Harvey & Gange (2003, b) have also provided data
for the duration of the first larval instar (about 5 months). Effects
of temperature, humidity and amount of nitrogen in the wood on development
are also unknown.
larvae coexist within a same stump (Paulian, 1959; GTLI, pers. obs.)
. However, any other details of larval demography are lacking: death
rate of each instar, predation or parasitism levels, within- or interspecific
competition, cannibalism (which has been reported for other xylophagous
beetles). Data presented by Murria Beltrán et al. (2004) are
the first confirmation of coexistence of larvae from L. cervus,
P. barbarossa and D. parallelepipedus in the same tree,
although details about spatial segregation within the tree are not provided.
After last larval molt, in which 10 cm length can be surpassed (Sánchez, 1983; GTLI, pers. obs.), pupation occurs in the soil, near the stump (de Ligondes, 1959; Sprecher, 2003; Harvey & Gange, 2003 b; GTLI, pers. obs. of specimens in terrarium). Pupation occurs within a chamber built with wood pieces, ground and other materials stuck together with saliva (de Ligondes, 1959; Español, 1973; Harvey & Gange, 2003 b; GTLI, pers. obs. of specimens in terrarium). Metamorphosis occurs during autumn and imagoes overwinter within the pupal chamber or in the ground nearby and show up at the end of next spring (de Ligondes, 1959; Rodríguez, 1989; Harvey & Gange, 2003 b; GTLI, pers. obs. of specimens in terrarium). The statement of Paulian (1959) that larvae overwinter before metamorphosis seems to be refuted by these more recent observations.
Larvae posses a stridulant organ. Sprecher (2003) has studied its morphology, but its function remains unknown.
(northwestern Spain), imagoes occur from middle June to the end of August
or early September, showing higher abundance during July and some between-year
variation (Álvarez Laó & Álvarez Laó,
1995). Phenological variation with altitude and latitude is also conceivable.
For example, in the United Kingdom imagoes are mainly observed between
May and August, with a maximum in June (Percy et al., 2000) and in Sweden,
flight period is June-July (Palm, 1959). Males appear a little earlier
than females (proterandry) (Percy et al., 2000; Sprecher, 2003; Harvey
& Gange, 2003 b; GTLI, pers. obs.). Abundance also shows between-year
variation (Paulian & Baraud, 1982). Four year cycles could be present
(Drake, 1994) although there is no quantitative study to support this
Crepuscular or nocturnal habits of imagoes have been traditionally noted (Paulian & Baraud, 1982) but it seems to be also some diurnal activity (Álvarez Laó & Álvarez Laó, 1995) that could be more important in Mediterranean areas (Lacroix, 1968; Arturo Baz, pers. comm.). Flight abilities seem, in principle, well developed. Drake (1994) states that only males flight regularly and Percy et al. (2000) also point out to a higher inclination to flight in males. Hallengren (1997) tried to tradiotrack some individuals but mostly failed. Sprecher-Uebersax (2001; Sprecher, 2003) utilised radiotracking to study adult dispersal. Males flough regularly and dispersed up to 200 m. Females never flough and showed shorter dispersal. There are XIX century tales about mass movements (Darwin, 1871; Lacroix, 1968; Paulian & Baraud, 1982). Anyway, atrophy of flight muscles after some time has been reported (Paulian, 1988), which could limit dispersal likelihood.
in several American Lucanid species has been studied (Mathieu, 1969)
but equivalent work about L. cervus is old and written in German,
which make it unreadable to us. Mating duration is disputed: short according
to Baraud (1993), a short mating or several mating episodes in a short
period according to Mathieu (1969), or lasting even several days according
to Huerta & Rodríguez (1988). Recent observations by Jason
Chapman et al. (unpubl. data) and GTLI's (pers. obs.) support the last
option, or al least a prolonged contact or escort by the male. Mating
duration is, probably, very variable and this prolongation could be
related to paternity insurance in a competitive environment. Several
studies with other insect species show that last male copulating with
a female fecundates most of her eggs (Eberhard et al., 1993). Some observations
indicate that males mate more than once (Hallengren, 1997; Sprecher,
2003). Harvey & Gange (2003 b) have found that females also mate
more than once.
a single batch of eggs (Harvey & Gange, 2003 b). Although it was
said that the eggs were laid in dead tree bark crevices, recent observations
by Sprecher (2003) and Harvey & Gange (2003 b) show that eggs are
laid in the soil, close to the dead wood, at about 25 cm depth. Percy
et al. (2000) list 34 different species as oviposition substrates. Females
individually lay (Huerta & Rodríguez, 1988; Harvey &
Gange, 2003 b) three to 20 eggs of large size (3 mm length; Baraud,
1993; Harvey & Gange, 2003 b).
offered a functional explanation to the obvious sexual dimorphism in
this beetle. Males fight to get the females, making of selective value
the development of the mandibles as weapons in such fights. Something
similar occurs not only in many Lucanids (Otte & Staiman, 1979)
but also in other Scarabaeoidea beetles (Palmer, 1978; Cook, 1987),
as well as in other insects and, of course, in mammals. In several species
of horned beetles an advantage of bigger males in getting
a mate has been reported (Palmer, 1978; Eberhard, 1979; Brown &
Bartalon, 1986; Siva-Jothy, 1987). Males with less developed weapons
use to lose fights and to die without mating. This is the basis for
the evolution of this trait. In the Stag Beetle, larger males seem to
win fights more often (Lagarde et al., 2005; Jason Chapman et al., unpubl.
The huge variation
in the development of the males mandibles has been studied by
many authors (Paulian, 1959; Lacroix, 1968,1969; Clark, 1977). These
studies showed that mandible size is related to body size and that there
is a gradual and continuous transition from smaller individuals with
small mandibles to bigger individuals with well developed mandibles.
Differences in larval feeding, related to nitrogen content in the decaying
wood from which larvae feed, can explain the variable final body size
of imagoes, but genetical factors could also been involved (Paulian,
1988). Interaction between genetic and environmental factors in the
production of dimorphism in horn length has been studied in Ontophagus
(Scarabaeidae) (Emlen, 1994, 1996, 1997; Moczek, 1998; Emlen & Nijout,
1999; Moczek & Emlen, 1999; Hunt & Simmons, 2000). Its generality
in other cases of dimorphism is yet to be shown.
In many Lucanidae
species (Arrow, 1939; Paulian, 1959; Otte & Stayman, 1979) two clearly
different forms of males have been found (this is the famous difference
between major and minor males). In these species, a logarithmic plot
of mandible size against body size does not gives a single straight
line but two separate ones. That means that both kinds of males obey
different growth rules and mandible size cannot be attributed to a mere
body size difference. Some other factor must be responsible. Eberhard
(1980) postulated a mechanism to explain these differences. First, differences
in substrate quality in which larvae develop must exist. This produces
body size differences between adult males. Small males lose more fights
and achieve a lower reproductive success. This selects for an alternative
mating behaviour in small males. Instead of fighting for females, they
sneak in the places in which females use to be. There they wait for
a chance and, while big males fight each other, small males reach females
and copulate with them. Incredible as this could sound, this alternative
behaviour is a very common phenomenon in species in which males fight
for females, from insects (Siva-Jothy, 1987) to fishes and amphibians
(Krebs & Davies, 1993).
What about Stag Beetle? Although presence of minor males has been accepted for a long time, studies quoted above did not find any support for two different growth patterns. However, Eberhard & Gutiérrez (1991) and Lagarde et al. (2005) found it, by using special statistical analyses and a large sample size. Our own data do not support such conclusion (Álvarez Laó et al., 1995). In our oppinion, this species is in a transition stage, not having developed a clear difference in alternative strategies. Our idea is based on Kawano (1989), who established the existence, within Lucanidae, of all transitional grades from species lacking sexual dimorphism to those with sexual dimorphism plus male polymorphism. Solution to this conundrum must wait to detailed studies of fight and mating behaviour in this species.
The status of
the species in Europe differ from country to country. It has gone extinct
in Denmark and Estonia. In the United Kingdom a decline is suspected
for the last decades (Clark, 1966; Percy et al., 2000; see, however,
Pratt, 2003, for a very critical view of this purported decline). Status
in Latvia is not optimistic (Dmitry Telnov, pers. comm.) and is critic
in Luxembourg (Marc Meyer, pers. comm.). In the Netherlands the species
does not seem to have declined from the 1980s (Cuppen, 1992). Jirí
Simandl (pers. comm.) states that it is common in lowlands in the Czech
Republic and Slovakia. In Spain, the Asociación española
de Entomología (Spanish Entomological Society) coordinated the
compilation of all available information about all the arthropods listed
in the Habitat Directive. GTLI collaborated in that task and, with the
help of lot of people, a first distribution map for Spain was obtained.
That map is the only currently available tool to assess the conservation
status of Stag Beelte in Spain. Based on that map, the threat level
of Stag Beetle is Spain was considered as LC (Least Concern) using the
This does not
mean that concern reasons are lacking. The main candidate is habitat
loss. Although usually this species has been considered to be dependent
on old oak woodlands (Quercus robur), in the Iberian peninsula
it is also present on other Quercus species, such as Q. pyrenaica
and Q. rotundifolia. In any case, its dependence on mature woodlands
is not clear either. In Asturias (northwestern Spain) it is present
in bocage areas, in which meadows are interspersed with small woodlands
and hedges. It occurs also in urban parks and Eucalyptus plantations,
suppossedly because of the presence of deciduous trees as Chesnut, Castanea
sativa, scattered within such plantations. In United Kingdom, this
species persists also in bocage and suburban habitats (Drake, 1994;
Percy et al., 2000). All this points to the fact that Stag Beetle is
quite tolerant to both habitat fragmentation and degradation, although
no study has tested this point. Probably, availability of dead wood
is more important than the kind or conservation status of the forest,
but this should be assessed quantitatively. An altitudinal limit around
600 m is often mentioned (Jirí Simandl, pers. comm.) but this
is plainly wrong, at least south from the Cantabrian range in Spain.
threath usually mentioned is collection (Sánchez, 1983; Huerta
& Rodríguez, 1988; Rodríguez, 1989). Despite not being
included in the CITES, trade of this species is forbidden in the United
Kingdom (Percy et al., 2000). It si difficult to assess the situation
in Spain. On one hand, SEPRONA (a branch of the Spanish police in charge
of nature protection) does not have any knowledge of illegal trade involving
Stag Beetle (José Delgado, pers. comm.). On the other hand, some
people has told us about Stag Beetle being sold in some petshops and
stamp collection shops. Frequency of these activities and the real impact
on natural populations are unknown. In insect conservation literature,
harvest is often considered little important (Pyle et al., 1981) as
a source of species extintion, even at a local level. In any case, we
could face a legal gap in this respect because Stag Beetle is not included
Finally, negative effects of pesticides or road casualties on Stag Beetle populations have not been studied in any detail.
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