Australian Hydraenid Beetles: Diversity, Ecology, Biogeography Nick Porch (ANU) & Phil Perkins (MCZ-Harvard)

Southeastern Australia Gymnochthebius Fossils

Victoria Northwest Crater 20,000 BP

Victoria Stony Creek Basin 1.8 Myr.

Distribution and BIOCLIM Predicted Distribution of Gymnochthebius

Northwest Crater

Distribution and BIOCLIM Predicted Distribution of Gymnochthebius clarki

Northwest Crater

Summary Bioclimatic Data for Gymnochthebius and G. clarki

Gymnochthebius G. clarki MIN MAX MIN MAX NWC Annual Mean Temperature 7.8 27.6 12.8 27.6 13.4 Max Temperature of Warmest Period 18.3 38.6 24.5 37.3 22.6 Min Temperature of Coldest Period -1.4 17.9 2.3 17.8 5.5 Temperature Annual Range (5-6) 15.7 33.6 16.4 30.6 17.1 Mean Temperature of Warmest Quarter 12.3 31.2 17.5 29.9 17 Mean Temperature of Coldest Quarter 3.7 24.8 7.9 24.8 9.8 Annual Precipitation 142 1763 338 1527 769 Precipitation of Warmest Quarter 32 736 46 677 120 Precipitation of Coldest Quarter 3 497 3 418 260

An Overview of the Family Hydraenidae

• Small to very small (c.0.7-3.5mm) • Worldwide (not some oceanic islands) • Forty world genera, est. 2400 species (1400 desc., 1000 new) • Larvae terrestrial or riparian • Adults generally aquatic-marginal aquatic – a few terrestrial • Non-swimming • Specialised ventral bubble for respiration • Most species fully winged, but poor dispersers • Very few taxonomic specialists – Perkins, Jäch

Taxonomic Work on Australian Hydraenidae

220 Ochthebius queenslandicus Hansen 200 2007 redescription 180

s 160 e i Perkins 2004-2007 revisions c e 140 172 new species p S

120 e v i 100 t a l Zwick 1977: Hydraena revision u 80 m u 60 Deane 1931-37 papers C 40 Hydraena luridipennis Macleay 1873 20 0 5 0 5 0 5 0 5 0 5 0 2 5 7 0 7 0 2 5 7 0 7 8 9 9 9 0 8 8 8 9

1 1 1 1 1 1 1 1 1 2 Year of Description Philip D. Perkins 2004-2007; 8 papers/monographs – c.840 pages

Classification and Affinities of Australian Hydraenidae

Ochthebiinae Ochthebiini Ochthebiina Gymnochthebius Orchymont 1943 Australia/NG & Americas Gymnanthelius Perkins Endemic Hughleechia Perkins 1981 Endemic Ochthebius Leach 1815 Cosmopolitan (SE Asia) Meropathina Tympanogaster Perkins 1979 Tympanogaster Perkins 1979 Endemic Topotympanogaster Perkins, 2006 Endemic Hygrotympanogaster Perkins, 2006 Endemic Plesiotympanogaster Perkins, 2006 Endemic Tympallopatrum Perkins 1997 Endemic Hydaeninae Hydraenini Hydraena Kegelann 1794 Cosmopolitan Limnebiini Limnebius Leach 1816 Cosmopolitan (South Africa) GENUS: Gymnanthelius Perkins

Reference: Perkins 2004a. A revision of the of the Australian endemic water beetle genus Gymnanthelius Perkins (Coleoptera: Hydraenidae). Zootaxa 585, 1-39.

• Distinguished from related Gymnochthebius easily by habitus

• Lateral hyaline border relatively equal both before and after mid-point of pronotum in contrast to Gymnochthebius where hind border is wider than fore border

Species Specimens Collections Distribution clypeatus (Deane, 1931) 718 27 BASSIAN-northern? cupreus (Deane, 1937) 52 14 BASSIAN heiroglyphicus (Deane, 1933) 41 2 BASSIAN lamingtonensis Perkins, 2004 4 1 BASSIAN-Lamington maxipunctatus Perkins, 2004 9 5 BASSIAN opacicollis Perkins, 2004 437 18 BASSIAN porchi Perkins, 2004 235 16 BASSIAN-Gippsland tunicus Perkins, 2004 247 4 BASSIAN

Habitat Types and Distribution

GROUP 1: Clean sand and gravel margin species clypeatus cupreus lamingtonensis heiroglyphicus

GROUP 2: Muddy margin species opacicollis tunicus porchi maxipunctatus?

Also now recorded for Carnarvon Range region & Atherton Tableland (single site).

Victoria: Tarra River, 1.5km S. Tarra Falls

2 species of Gymnanthelius tunicus & porchi - 84 specimens also Gymnochthebius (1 species) 12 specimens Traralgon Creek: Type Locality Gymnanthelius tunicus Gymnanthelius opacicollis

McKillops Bridge area, Snowy River Gymnanthelius clypeatus Gymnanthelius cupreus GENUS: Gymnochthebius Orchymont

• Distinguished from related Gymnanthelius easily by habitus – pronotum not cordate • Lateral hyaline hind border is wider than fore border • 33 endemic Australian species, 2 in PNG, 1 shared; c.25 species North & South America • Many widespread species, but also significant number of range restricted species. e.g. artesian spring species • Moderately well-known; 11 from single localities although includes some NREs

Revision: Perkins, P.D. (2005). A revision of the water beetle genus Gymnochthebius Orchymont (Coleoptera: Hydraenidae) for Australia and Papua New Guinea. Zootaxa 1024, 1-161. Identifying Australia Gymnochthebius: Key Characters

Dorsal Structure of Pronotum is extremely variable: shape and size development median groove and lateral fovea microsulpture nature and extent size and shape of dorsal setae

Identifying Australia Gymnochthebius: Key Characters

Ventral structure and amount of hydrofuge pubescence: midlongitudinal carina on proternum present or absent metasternal glabrous areas – shape, size and microsculture extent of hydrofuge pubesence on abdominal ventrites Elytral shape, elytral setae and striation: presence or absence of arcuate setal fringe on elytral margin serial versus confused elytral striae shape of elytron: short and broad versus elongate convexity of elytron dorsal punctures and setation nature of elytral striae and intervals Habitus, microsculpture and setation: overall colour microsulpture extent and nature density, extent and nature of setae

Gymnochthebius: Ecology and Biogeography

• Generally marginal aquatic on muddy-sandy substrates

• Can be exceedingly abundant – 100s per metre of habitat

• Several species very tolerant of human modified waterscapes – others less so

• Range from dense Mountain Ash forest/Rainforest streams to arid zone freshwater and springs

Gymnochthebius: Predicted Species Richness

GENUS: Ochthebius Leach

• Single - potentially introduced - species in Australia. Closest relatives in SE Asia (Thailand, Sumatra)

• Widespread with records from major northern and western Australian port regions

• Probably saline and coastal wetland inhabitant

Revision: Perkins, P.D. 2007c. A review of the coastal marsh water beetle Ochthebius queenslandicus Hansen (Coleoptera: Hydraenidae). Zootaxa 1625, 35–42.

Ochthebius Darwin queenslandicus yellow pentagons

Normanton Townsville

Perth

GENUS: Tympallopatrum Perkins

• Southwestern Australia endemic genus • Four described species • Genus is terrestrial; extremely reduced hydrofuge pubescence • Found in wet-dry forests of southern Southwestern Australia Revision: Perkins, P.D. 2004b. A revision of the Western Australian endemic humicolous beetle genus Tympallopatrum Perkins (Coleoptera: Hydraenidae). Zootaxa, 672, 1-16.

MAP (mm) 150-250 250-500 500-750 750-1000 1000-1250 >1250

")") ") ") ") ") ") ")") ") ") ") ") Tympallopatrum ") GENUS: Hughleechia Perkins

• Australian endemic genus • Two described species: • Southern Australia H. giulianii (left) • Tasmania H. gracilis (right) • Genus marginal marine rocky habitats including splash pools, crevices and supra- tidal rock-faces Revision: Perkins, P.D. 2007b. A revision of the Australian intertidal water beetle genus Hughleechia Perkins (Coleoptera: Hydraenidae). Zootaxa, 1527, 17-29.

Green Cape: habitat of H. guilianii

GENUS: Limnebius Leach

• Australian/NG species recognised by habitus • Closest relative is South African • Widespread - often locally common - in a variety of mud-sand based substrates especially beside large slow rivers • Absent from Western Australia and Tasmania – predicted marginally suitable • Exhibits marked clinal variation in aedeagus morphology Revision: Perkins, P.D. 2004c. Limnebius acupunctus, a new species of water beetle from Australia and Papua New Guinea (Coleoptera: Hydraenidae). Zootaxa, 749, 1-12.

Limnebius: bioclimatically predicted distribution

GENUS: Tympanogaster Perkins

• Australian endemic genus with four subgenera • Hygrotympanogaster, Tympanogaster, Plesiotympanogaster, Topotympanogaster

• Strictly freshwater (Hygros, Tymps, Plesios) to terrestrial (Topos)

Revision: Perkins, P.D. 2006. A revision of the Australian humicolous and hygropetric water beetle genus Tympanogaster Perkins, and comparative morphology of the Meropathina (Coleoptera : Hydraenidae). Zootaxa 1346, 1-396.

Tympanogaster Pleisiomorphic subgenera and linking taxa

Apomorphic Tympanogaster s.l. larvae • Larvae common in same habitat as adults • Can occur at extremely high densities with adults • meso-thoracic spiracles are dorsally oriented and tube- like for respriation in hygropetric habitats (larvae only known for Tymps and Hygros

lateral Tymp dorsal

Hygro Identifying Tympanogaster: Reality Bites

Perkins 2006 states “Tympanogaster identification can be difficult” and “As in many other members of the family, detailed study of the male genitalia is usually essential for reliable identifications”.

Geography is a very reliable character given the high degree of narrow range endemism, however, until this genus is well collected there will remain questions about newly collected material. Even where well collected – like the Grampians – there are taxa know from single sites.

subgenus: Tympanogaster Perkins

• 38 described species but many more awaiting discovery and description

• Most very similar looking

• Long legs, stout claws and distinctive habitus optimal for fast water streams

• Splash zone of boulders in fast steams or less commonly, rockfaces in streams

• No polymorphic males (cf. Hygro- tympanogaster, below)

• High levels of narrow range endemism

Talbot Creek, Baw Baw Range habitat of T. modulatrix

Water Shute habitat of Tympanogaster

Williams River, Barrington Tops ‘Classic’ habitat of Tympanogaster Williams River, Barrington Tops subgenus: Hygtotympanogaster Perkins

• 37 described species but many more awaiting discovery and description

• Generally hygropetric on rock adjacent toflowing water; in seeps, on rockfaces, waterfalls etc.

• Males frequently polymorphic in better collected species but this may not always be so

• High levels of narrow range and extreme (single site) narrow range endemism

• Almost exclusively (one exception) restricted to eastern Australian forest habitats

Allometric Polymorphism in Hygrotympanogaster & ‘hygropetric combat’

• Large males have markedly enlarge labral processes • Males of polymorphic species have stout peg-like processes on their mesotarsi • Polymorphic species are hygropetric

• Males fight: Perkins speculates that they fight for prime realestate and therefore food and female

H. novicia female (middle)

male morphs (outside) River margin perennial streams and associates seeps

Type locality for H. atroargenata Perkins H. megamorpha Perkins

Late 19th early 20th Century H. bondi Perkins Bondi Heights Tamarama H. wahroonga Perkins

Perennial seeps over rock Marginal rockface flows

Splash zones of waterfalls and streams

subgenus: Topotympanogaster Perkins

• 8 species known from 11 sites in the Wet Tropics region of north Queensland

• Only two species known from more than one collection event suggesting many more remain to be discovered

• Thought to be largely humicolous relative to other Tympanogaster subgenera although exceptions

• Species keyed predominantly using distinctive pronotal characters

Latitudinal Diversity of Typmpanogaster sensu lato

0% 20% 40% 60% 80% 100%

Wet Tropics (22) Topotympanogaster N

Border Range Region (11)

NewEngland/Dorrigo (14) Tympanogaster s.s.

BarringtonTops Region (5)

Sydney/BlueMtns/Canobolas (9) Hygrotympanogaster

South of Sydney (mainland) (19)

Tasmania (3) Pleisotympanogaster S

Patterns of Endemism Tympanogaster s.s. (main) Topotympanogaster (inset) Plesiotympanogaster (inset)

Patterns of Endemism Hygrotympanogaster

Undescribed

subgenus: Hydraena Kegelann

• 84 described species in 24 species groups

• Species group richest in mesic zones with high levels of regional endemism

• One species group (Rudallensis sp. grp.) with number of widespread arid zone taxa

• Dominantly found in the gravelly/sandy margins of streams – limited number of still water species. One terrestrial species (hamifera)

Revision: Perkins, P.D. 2007a. A revision of the Australian species of the water beetle genus Hydraena Kugelann (Coleoptera: Hydraenidae). Zootaxa 1489, 1-207.

Identifying Hydraena species groups: Elytral and Habitus

• Body size frequently relatively constant in species group • Microsculpture arrangement and nature • Colour pattern; uniformity, pattern, maculae • Shape and size of elytra; relative to pronotum; punctuation/striation

Identifying Hydraena species groups: Pronotal Characters • Australian Hydraena have up to four pairs of pronotal foveae • Their presence/absence, depth, and arrangement constant in sp. grps. • Pronotal shape and relative measurements relatively uniform • Surface sculpture; punctuation, microsculpture

Identifying Hydraena species groups: Metaventral plaques

• Two metaventral plaques – areas without hydrofuge pubescence • Plaques vary in shape – relatively constant within species groups

l - plaque length s – plaque separation w – plaque width p2 – mesoventral intercoxal process separation

Identifying Hydraena species groups: Female last tergite X

• Shape of last tergite X constant within species groups • Spcifically, number of impressions

No impression Single impression Three impressions Reticuloides Barbipes Affirminata Billi Ambiflagellata Bidefensa Cultrata Castanea Cunninghamensis Invicta Hamifera Luridipennis Queenslandica Magnetica Rudallenis Porchi Textila Simplicicollis Trapezoidalis Weiri Tridisca Ypsilon Wattsi Zwicki Identifying Hydraena species: Male Genitalia

• As with many hydraenids the male genitalia are highly diagnostic and must be studied for reliable species determination

• This is unlikely to change with future work although detailed morphometric work and interactive keys may, in the future, enable species level identifications

• Hydraena genitalia are complex and asymmetric

Hydraena Castanea grp. Hydraena antaria Perkins, 2007 187 7 BASSIAN Hydraena castanea Deane, 1937 657 47 BASSIAN Hydraena clavigera Zwick, 191 66 34 WIDE-Dividing Range Hydraena dorrigoensis Perkins, 2007 1 1 DORRIGO-NRE Hydraena luminicollis Perkins, 2007 189 7 sQLD/nNSW MTNS Hydraena spinissima Perkins, 2007 4 4 sQLD/nNSW MTNS Hydraena tricantha Zwick, 197 148 38 BASSIAN Hydraena Ambiflagellata grp. Hydraena ambiflagellata Zwick, 1977 77 18 BASSIAN Hydraena cygnus Zwick, 1977 5 2 SOUTHWESTERN Hydraena decipiens Zwick, 1977 4 2 WIDE Hydraena Textila grp. Hydraena ascensa Perkins, 2007 83 7 WET TROPICS Hydraena capetribensis Perkins, 2007 1 1 WET TROPICS Hydraena finniganensis Perkins, 2007 1 1 WET TROPICS Hydraena intraangulata Perkins, 2007 20 6 TORRESIAN-Cape York Hydraena storeyi Perkins, 2007 7 4 WET TROPICS Hydraena textile Perkins, 2007 1 1 WET TROPICS Hydraena Billi grp. Hydraena biimpressa Perkins, 2007 1 1 WET TROPICS-NRE Hydraena billi Zwick, 1977 7 9 BASSIAN-Central Highlands Hydraena hynesi Zwick, 1977 158 23 BASSIAN Hydraena monteithi Perkins, 2007 1 1 WET TROPICS-NRE Hydraena Rudallensis grp. Rudallensis Group Hydraena ambiosina Perkins, 2007 46 6 TORRESIAN/TIMORIAN Hydraena athertonica Perkins, 2007 12 4 WET TROPICS Hydraena australica Zwick, 1977 279 51 TORRESIAN/TIMORIAN Hydraena australula Perkins, 2007 2 2 Kimberley Hydraena blackburni Zaitzeff, 1908 160 38 TORRESIAN/TIMORIAN Hydraena ferethula Perkins, 2007 1 1 TIMORIAN Hydraena kakadu Perkins, 2007 2 2 TIMORIAN Hydraena lucernae Zwick, 1977 106 32 TORRESIAN/TIMORIAN Hydraena parciplumea Perkins, 2007 1 1 TIMORIAN Hydraena parva Zwick, 1977 29 10 TORRESIAN/TIMORIAN Hydraena rudallensis Blackburn, 1896 218 60 WIDE Hydraena sp. grps.: Predicted Species Richness

Hydraena sp. grps.: Predicted Species Richness

Wet Tropics