A Guide to Stag Beetles of Australia

Part I

Classification

An important foundation stone of human thinking is the urge to sort things. Every time when we come face to face with something unfamiliar, we try to fit it into some kind of a system in order to understand what it is.

Scientific classification (the science of taxonomy or biological systematics) is a method by which biologists group and categorise extinct and living organisms. Modern classification is based on the system of Carolus Linnaeus, the 18th century Swedish scientist.

This system has been revised several times in order to accommodate the results of new research, especially the Darwinian principle of evolution and more recently genomic analysis through DNA sequencing.

A perfect system of classification has yet to be developed. What we have – the modern version of the Linnaean system – needs constant honing and this is what taxonomists do all around the world.

The ingenuity of the Linnaean system is manifested by its flexibility and by its ability to accept changes without getting destroyed in the process. Ever since its birth, many new species are being added in the world taxonomic ‘inventory’ each year. Now more than 1.5 million species of organisms bear names, given to them according to the Linnaean system.

Apart from the fact that the Linnaean system of classification is capable of accommodating an endless deluge of new species, it has another great feature – binominal nomenclature.

Linnaeus insisted that every species shall have a Latin name, consisting of two words. This method of creating names in this way is known as binomial nomenclature.

The Linnaean binomial nomenclature is simple and practical. Every species has a ‘surname’, known as generic name, which signifies that it belongs to a particular group known as a genus (plural: genera), and a ‘given name’, known as the specific name, which denotes its identity as a species. The combination of these two names is the scientific name of a species. There could be many species within one genus and in that case they all shall have the same generic name, but their specific names must all be different. No two or more species of any organism should have the same generic and specific name in one combination. However, a specific name can be given to more than one species, providing they all belong to different genera.

A scientific name in reality consists of more than two words. It is customary to give the generic and specific names first, followed by the name of the person, known as the author, who described and named the species. After that, a four-digit number may be placed, indicating the year when the name was created. For instance Lucanus cervus Linnaeus, 1758 means that the species cervus belongs to the genus Lucanus, and it was named by Linnaeus in 1758. When the author’s name is in parenthesis: Geotrupes stercorarius (Linnaeus, 1758), it means that the species was initially named by him but later on it was transferred from its original genus to another. Linnaeus initially described and named this species as Scarabaeus stercorarius, but later on, when the genus Geotrupes was erected, stercorarius was transferred to it.

When a species is renamed, the old name becomes a synonym. In scientific works, these synonyms are usually mentioned in addition to the valid names in order to avoid confusion.

Traditionally, generic and specific names are printed in italics and in handwriting or, if written by a typewriter, without italicised letters and underlined.

Since the mid-1980s DNA analysis became more and more important in identifying species in entomology. The process is quite complex, relatively expensive and slow if compared with the traditional entomological process for determining species by their morphological characteristics.

At present, DNA analysis is not within the reach of most amateurs and only a certain percentage of professional researchers have frequent access to genome-investigating methods and equipment. But it is more than likely that in time techniques will improve and become more accessible to a greater number of entomologists. If this becomes a reality, taxonomy will undergo tremendous changes, many species will be reassigned, higher taxa will be revised and the current face of entomology will change forever.

The language of entomology

Ancient Greek and especially Latin used to be the languages of science in Europe. The teachings of the earliest Greek and Roman scientists were respected and accepted throughout millennia. Time, however, didn’t stop and the world kept changing; the ancient Greeks and Romans vanished and their languages would have died with them, if scientists and the practitioners of Christian religions didn’t use them. Why did they keep using these ‘dead’ languages? A ‘dead’ language is not subject to changes, so a script made in Latin or Ancient Greek means the same today as the day it was written – even if it was 2000 years ago. This would be impossible to achieve using a ‘living’ language, which could change within a relatively short time, so much so that it would become unintelligible or, at best, difficult to understand precisely. More than likely, this was the main reason why early naturalists, such as Linnaeus, chose classical Latin to describe and name newly discovered species of plants and animals.

As methods of research advanced, more and more detailed and precise descriptions were required. Pure Latin, being a ‘dead’ language, soon became inadequate for the task. It became very difficult to speak and write Latin fluently in a continuous and expressive manner, and more and more frequently scientists resorted to the most widely used spoken and written European languages. By the 19th century German, French and English became the languages of scientific publications aimed at international audiences. However, Latin and Greek words were used when these languages couldn’t describe particular features or habits of an organism in a concise and precise manner. By using the grammar and vocabulary of a ‘live’ language but adding as many Latin and Greek words as necessary, a scientific language developed. Workers of many nationalities followed this method to create their own versions of a scientific language, based on their mother tongues. Difficulties in communication between the scientific communities of various nations led to the realisation that one language should eventually dominate science worldwide. In the late 20th and early 21st centuries, the English language became the dominant international language. It is not only customary but also practical to publish scientific works in this language, as it is now widely understood all over the world. Publishers and authors often find it necessary to include English summaries of their non-English-language publications.

When an organism and its features are described in a scientific manner, the description must be as accurate as possible. The best way to do this is to avoid expressions that could be perceived differently by various readers and instead use accurate, purposeful words in order to convey a concise description. No living language has a vocabulary that would be suitable for this job without blending some Latin and/or Ancient Greek into it.

Morphology

The scarab-like beetles:

Scarabaeoidea

The stag beetles, the Lucanidae, are a family within the superfamily Scarabaeoidea. Members of this superfamily generally have lamellate antennae. The terminal segments of this highly developed organ form a club, composed of plates known as lamellae. The typical scarab lamellae can be compressed into a ball or spread out in a fan-like manner. In the Lucanidae the antennal club is quite variable and is in some species flabellate rather than lamellate.

The main morphological characters of the Australian Lucanidae

Body elongate, some robust, 5–72 mm

Head prognathous, males often with prominent mandibles, some branched

Antennae with long scape, often geniculate, three to seven loose segmented club that cannot be completely closed

Eyes entire or with a partial or complete canthus

Labrum and clypeus usually fused to frons

Scutellum visible

Abdomen with five visible ventrites

Legs relatively long and slender, protibia well developed

Tarsal formula 5–5–5

Usually brown or black, some species have lighter dorsal patterns, some vividly metallic.

An important character that is used to differentiate lucanids from other scarabs is the number of segments in the club. In the Australian Lucanidae the antennal club is composed of three to seven segments and the antennae have 10 segments in total. The antennae vary from being geniculate (elbowed) to non-geniculate and have a relatively long scape (first segment of the antenna). The genus Lissapterus has at the most only a slightly pectinate antennal club whereas in the genus Ceratognathus the comb-like segments are quite long (especially in males) and movable, but cannot be closed fully into a compact club.

A very conspicuous character of these beetles is their sexual dimorphism. Males of many species have large, ornate mandibles, while females have smaller and very simple ones. The mandibles of male Lucanidae are often used in combat with rival males, especially when attempting to mate with a nearby female or in self-defence against would-be predators. Usually it is quite easy to distinguish the sexes through their mandibles, although in some cases the smallest males look very much like females and several species show only minimal sexual dimorphism. The sexes are indistinguishable externally in some genera (e.g. Figulus).

Another well-known character in this family is the allometric development of the male mandible (the size of the mandible is proportional to the size of the body), which has often confounded taxonomists. The marked differences in the teeth of the mandibles may depend on the overall body size of the male. In this book, where this occurs, we will simply refer to small males of a species as ‘minor males’ and large males as ‘major males’.

Schematic map of a non-specific stag beetle (based on Ryssonotus nebulosus), dorsal view. 1. Frons, 2. Pronotum, 2a. Disc, 3. Scutellum, 4. Suture, 5. Elytra, 6. Hind-or metafemur, 7. Hind- or metatarsus, 8. Hind- or metatibia, 9. Mesotarsus, 10. Mesotibia, 11. Mesofemur, 12. Mesothorax, 13. Fore- or profemur, 14. Fore- or protibia, 15. Fore- or protarsus, 16. Maxillary palp, 17. Antenna, 18. Mandible, 19. Eye, 20. Vertex. Drawing: Esther Bolz.

Schematic map of the head of a non-specific stag beetle (based on Lucaninae female), dorsal view. 1. Frons, 2. Vertex, 3. Mandible, 4. Labrum, 5. Canthus, 6. Eye, 7. Postocular margin, 8. Preocular margin, 9. Galea (outer lobe of the maxilla). Drawing: Esther Bolz, after Holloway.

Schematic map of the head of a non-specific stag beetle (based on Lucaninae female), ventral view. 1. Labrum, 2. Mentum, 3. Gula (throat), 4. Gena (cheek), 5. Eye, 6. Canthus, 7. Antenna, 8. Galea (outer lobe of the maxilla), 9. Mandible, 10. Maxillary palp, 11. Labial palp. Drawing: Esther Bolz after Holloway.

Antenna of Syndesus cornutus. Drawing: Esther Bolz.

Antenna of a male Ceratognathus sp. Drawing: Esther Bolz.

Antenna of a Lamprima sp. Drawing: Esther Bolz.

Australian lucanids are mostly black, dark brown or, in the case of the subfamily Lampriminae, a vivid colour with a metallic lustre. The scutellum is always visible, although sometimes it may be rather small.

The Lucanidae

Stag beetles have always interested coleopterists, amateurs and professionals alike. Many of these beetles have spectacular shapes, their larvae lead cryptic lives and, in most species, people do not commonly encounter the adults in their habitats. The Lampriminae and many of the tropical Lucaninae are splendidly coloured while others show an amazing variety in the male mandibular size and structure – these are some of the outstanding characteristics of this family which made these beetles so popular. Their aesthetic appeal and the rarity of some species represent great value to collectors. In several cultures the stag beetle represents a mythical or mystical element and some important representations of it can be found already in Renaissance art (e.g. Dürer’s famous drawing of the European stag beetle). Undoubtedly, the study of this beetle family has great scientific importance.

The family Lucanidae consists of scarab type beetles belonging to the superfamily Scarabaeoidea. In the past this family was considered to be the most primitive family within the Scarabaeoidea (Lawrence and Newton 1995). More recent research, however, proposes that the Glaresidae (glaresid beetles) are the most primitive family of scarabs and places the Lucanidae between Glaresidae and Trogidae (skin or hide beetles) and in the same clade with Passalidae (bess beetles) and Bolboceratinae (Geotrupidae, earth-boring dung beetles) (Smith et al. 2006). Glaresidae are small, 2.5–6.5 mm long beetles that don’t occur in Australia and practically nothing is known about their biology.

Some lucanid beetles have secluded, long life cycles, spent mostly inside or under decaying, fungus-ridden logs or standing dead and rotting trees. Development from egg to adult can take several years, depending on the species and local circumstances. Lucanid larvae, like most other wood-eating insects, must employ the help of microorganisms to digest food with high cellulose content. Recent studies have shown that female lucanid beetles possess microbe-storing organs (mycangia) containing microorganisms closely related to xylose-fermenting yeasts (Tanahashi et al. 2010). After laying an egg, the female discharges some of these microorganisms on and around it, thus leaving a digestion aiding ‘starter pack’ for the offspring (Fremlin 2015). So far this research has focused only on European and Japanese lucanid species – the Australian lucanid fauna has yet to be investigated.

As the developing larva grows, it moults several times. The stages between moults are known as instars. Stag beetle larvae have three instars. Once the larva reaches its full size, it pupates. For pupation to happen, the larva must first make a cavity (pupal chamber) within the rotting wood or in the soil where it developed. Pupal duration is usually several weeks or a month depending on the species, and the newly emerged adult needs to harden and will remain in the pupal chamber for a similar amount of time or for many months if the time of year or conditions are not appropriate. Young beetles with unhardened exoskeletons and immature colouring are known as teneral adults.

The lucanid larva has typical scarabaeoid characteristics as it appears as the usual, whitish, C-shaped scarab ‘curl grub’. The head is strongly sclerotised and the mandibles are stout and strong, adapted to feed on decaying wood.

Lucanid larvae can stridulate, although this may not be readily observed and is often not audible to the human ear. It is thought that stridulation helps to establish the presence of a larva to other larvae in a population and may help in promoting avoidance behaviour. Lucanid larvae can be distinguished from the larvae of other scarab families by the location of the stridulatory apparatus on the mid and hind coxae of the legs and by the large fleshy pads on either side of the vertical or Y-shaped anal opening.

Stag beetles have exarate pupae. The appendages are free and quite recognisable, but movement is limited to the abdominal segments. The pupa may wriggle a little when irritated, but otherwise remains inactive.

While larvae consume considerable amounts of rotten wood and have even been found in rotten structural timber (Lawrence 1981a), it is believed that adults of many species eat very little or don’t eat at all. This assumption is based on the study of some species’ reduced mouthparts and studies of their alimentary canal. Adults of some Lamprima species feed on plant secretions and possibly nectar and certainly do feed on soft fruit when kept in captivity. Other species (e.g. Cacostomus squamosus) are known to feed on the exuding sap of injured trees.

For a large percentage of Australian Lucanidae, nothing much is known about adult feeding nor breeding habits. The biology of the Australian stag beetles is not thoroughly researched although some species are successfully bred in captivity. This topic is discussed with more details on pages 214–217.

A characteristic stag beetle larva. Photo: PZ.

The sclerotised head of a stag beetle larva. Photo: PZ.

The large fleshy pads at the posterior of a stag beetle larva. Photo: GH.

Origins of the Australian stag beetle fauna

The Australian fauna shows definitive connections to the fauna of the ancient Gondwana and also to Asia. The adjective ‘Gondwanan’ is often used in biogeography, the study of the distribution of species and ecosystems in geographic space and through geological time. Gondwana, the ancient, enormous continent, included most of the landmasses in today’s Southern Hemisphere, including Antarctica, South America, Australia and also some others that are now north of the Equator. This supercontinent started to break up in the early Jurassic, ~180 million years ago. The parts that separated moved apart, taking with them their flora and fauna, like travellers on gigantic rafts, dispersing over huge areas. Many aspects of Southern Hemisphere biogeography refer to patterns of distribution of living organisms, especially when these seem to be related and are restricted to two or more of the now discontinuous regions that were once part of Gondwana.

A stag beetle that can fly: Lamprima latreillii Macleay, 1819. Photo: PZ.

The discovery of two extant species related to Sphaenognathus (p. 73) and one extinct, fossil Syndesus (p. 68) species are perhaps the best examples (in Lucanidae) of our Gondwanan origins.

Biogeographic connections between Australia and Asia are still poorly understood, although the plate tectonics of the Indo-Pacific region is now well described. During the Pleistocene (2 588 000 to 11 700 years ago), extended periods of glaciation caused decreases of sea levels by more than 100 m in Australasia. During these periods, landbridges formed between South-East Asia and northern Australia. Migration of plants and animals was possible throughout the epoch.

Ancestors of some of our stag beetle species belonging to genera Dorculus, Dorcus, Prosopocoilus and Figulus could have arrived from Asia through the connecting expanses of dry land.

Some of our endemic species are good fliers but others, mainly those that exist in much more specialised habitats and lead cryptic lives, have lost the ability to fly; examples are Lissapterus and Lissotes, which live under or within decaying timber.

Kim and Farrell (2015) recently researched lucanid relationships and divergence time estimates and how Gondwanan break-up effected biogeographical developments.

Taxonomic studies of the Australian stag beetle fauna

The description of a Tasmanian stag beetle, Lucanus cancroides (known now as Lissotes cancroides), was among the first Australian beetles described by the Danish entomologist Johan Christian Fabricius, in 1787 (Fabricius 1787). In the following years, more and more insect specimens were brought back to Europe and entomologists, following Fabricius, described several new species from the newly explored Great Southern Land that later became known as Australia.

Pierre André Latreille, the ‘Prince of Entomologists’, was the first entomologist to focus on the uniqueness of the Australian fauna and described the first Australian genus Lamprima in 1807 (Latreille 1807), recognising the Norfolk Island species (Lamprima aenea) as its type species. In 1817 he described an additional species, Lamprima aurata. William Sharp Macleay in 1819 described the endemic genus Ryssonotus (Macleay 1819). In the same year he also described the more cosmopolitan genera of Aegus and Figulus (Macleay 1819), which were later recognised by other researchers to have some Australian endemic species. Another entomologist, Edward Newman, described the genus Cacostomus in 1840 (Newman 1840).

The endemic genus Lissapterus was first recognised and described by Henri Deyrolle in 1870 (Deyrolle 1870), and in 1881 he described several species in the genus Lissotes (Deyrolle 1881) and one species in the genus Ryssonotus (Deyrolle 1881). Boileau, Didier and other French entomologists followed in their footsteps and named more species, while their German colleagues H. Burmeister and G. Albers also got interested and described at least four newly discovered Australian stag beetles.

The most prolific British worker of the time was J. O. Westwood, who described in total 19 currently recognised Australian species and three new genera, two of which are endemic to Australia (Ceratognathus and Lissotes) (see the references in the species descriptions). On several occasions this work was done in collaboration with F. W. Hope.

Another British entomologist F. J. S. Parry described Pseudodorcus in 1870 (Parry 1870) and the Tasmanian genus Hoplogonus in 1875.

As the new settlers began to develop and explore Australia, scientific investigations uncovered more interesting species. The first major Australian entomologist was William John Macleay. He was born in Scotland and came as an 18-year-old lad to Australia in 1839. However, before he actively engaged in scientific endeavours, he became a pastoralist and a politician. In 1857 he married and from then on devoted much of his time and energies to zoology. He has described two Australian stag beetle genera: Homolamprima, including one species, Homolamprima crenulata (Macleay 1885), and Phalacrognathus, also including one species, our ‘legendary’ Phalacrognathus muelleri (Macleay 1885). He also formed an entomological society in Sydney and later became the president of the Linnaean Society of New South Wales. His achievements in politics and science earned him a knighthood.

The English-born Australian Thomas Blackburn was an ordained priest of the Church of England. He had a very keen interest in entomology and became most active towards the end of the 19th and the beginning of the 20th centuries. Blackburn described 3069 Australian invertebrate species, mostly beetles, including three stag beetles in the genus Ceratognathus (see the reference in the species descriptions).

In the first half of the 20th century the rate of discoveries slowed a little, but this didn’t mean that the Australian stag beetle fauna had been thoroughly explored. While it seemed that the bulk of the stag beetle species were already named, this assumption was soon proven wrong as a ‘new breed’ of Australian researchers, A. M. Lea and H. J. Carter, soon described many new species. This probably provided an impetus that intensified interest among overseas specialists. The Slovenian-born American entomologist Bernard Benesh named another two species in 1943. In 1953, the Frenchmen R. Didier and E. Seguy published a catalogue listing most of the known Lucanidae of the World. This multi-volume work is beautifully illustrated with the graphic habitus portraits of many species by L. M. Planet (Didier and Séguy 1953). Benesh also produced the World Catalogue of Lucanidae in 1960, listing ~1500 species – among them all the then-known Australian species (Benesh 1960). The French Lucanidae specialist Melchior de Lisle described Figulus howei (de Lisle 1967) from Lord Howe Island in 1967.

The discovery, description and naming of new species are important elements of entomological research, but there are quite a few eminent entomologists who have not focused on describing new species, but rather focused their efforts to ‘clean up’ the higher levels of taxonomy. One such Entomologist was B. A. Holloway (New Zealand), who in the 1960s carried out detailed comparative morphological research that led her to refine the higher classification of the stag beetles (Holloway 1960).

While the majority of coleopterists study mostly the adults of a species, larval research also has great importance. The first in-depth studies of the larvae of the Lucanidae were carried out by F. I. van Emden in 1935 (van Emden 1935) and by J. F. Lawrence in 1981 (Lawrence 1981b). In this paper Lawrence produced a larval key to some of the major groups of stag beetles, largely based on the key of Emden.

The discovery of a Sphaenognathus- like species in Queensland was a great entomological sensation. Until this discovery Sphaenognathus was only known from the Neotropical region (South America) and the Australian species was perceived as a faunal remnant of Gondwanan times. The male of this species was described and named Sphaenognathus queenslandicus by B. P. Moore in 1978 (Moore 1978). In the late 1980s, Belgian-born Hugues E. Bomans studied the Australian fauna very thoroughly and discovered 11 new species in the Lucaninae subfamily. His work was greatly helped by the collaboration of Hungarian-born Australian coleopterist George F. Bornemissza and several young entomologists who carried out extensive field studies of the stag beetles in Tasmania. Luca Bartolozzi (Italy) has worked on and still works on the Australian stag beetle fauna. He has described new Hoplogonus and Lissotes species from Tasmania during the last 20 years