Abstract
New Zealand has a diverse and highly endemic fauna of dance flies (Empididae) with much of its diversity found in the tribe Hilarini. Here we report the first fossil of a dance fly from New Zealand from lacustrine diatomites of the earliest Miocene Foulden Maar Fossil-Lagerstätte in Otago. Hilarempis otagoensis sp. nov. is described from a single male specimen and assigned to the tribe Hilarini based on characters of the wing venation. This new species represents the first fossil record of the predominantly southern temperate genus Hilarempis Bezzi and the second fossil record of the subfamily Empidinae from the Southern Hemisphere. The articulated preservation suggests that Hilarempis otagoensis sp. nov. lived among riparian vegetation on the shore of the Foulden Maar palaeolake and probably drowned while skimming the water surface in search for prey or nuptial gifts.
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Introduction
The Empidoidea (Diptera) include more than 12.000 species of mostly predatory flies that are grouped into five to nine families and numerous subfamilies (Sinclair and Cumming 2006; Pape et al. 2011; Wahlberg and Johanson 2018). The fossil record indicates that empidoid flies existed in the Jurassic and that families and subfamilies diversified during the Early Cretaceous (Grimaldi 1999, Grimaldi and Cumming 1999, Grimaldi and Engel 2005). The Empididae (dance flies) is divided into several subfamilies, including the Clinocerinae, Empidinae and Hemerodromiinae (Sinclair and Cumming 2006). About 55 fossil species of Empidinae have been described, mainly from Cenozoic sites of the Northern Hemisphere, and most are placed in the extant genera Rhamphomyia Meigen and Empis Linnaeus of the tribe Empidini (e.g. Statz 1940; Förster 1891; Evenhuis 1994). The oldest and only Mesozoic records of Empidinae are Turonempis styx Grimaldi and Cumming and Emplita casei Grimaldi and Cumming (tribe Empidini) from New Jersey amber (Turonian) and Empis orapaensis Waters (tribe Empidini) from the Cenomanian/Turonian of Botswana, with the latter species representing the sole Southern Hemisphere fossil record of Empidinae (Grimaldi and Cumming 1999, Waters 1989). Fossils of the tribe Hilarini are extremely rare and exclusively represented by species of Hilara Meigen from the Palearctic Region (e.g. Meunier 1908, 1915; Gil Collado 1926; Theobald 1937; Evenhuis 1994).
Here we describe a fossil specimen of Hilarempis Bezzi, a hilarine genus with 110 extant species that occur mainly in southern temperate areas (Rafael 2012). The specimen comes from the Foulden Maar Fossil-Lagerstätte in southern New Zealand. The geographically isolated islands of New Zealand exhibit a diverse extant Empidoidea (exclusive of Dolichopodidae) fauna of 41 genera and at least 300 (plus many undescribed) species, with all of the described species being endemic to the islands (MacFarlane et al. 2010). Much of their overall diversity, life history and ecology remain to be studied in detail (MacFarlane et al. 2010, and references therein). The tribe Hilarini (Empidinae) is particularly well represented in New Zealand (47 species in 6 genera) and the most speciose genera are Hilara (15 spp.) and Hilarempis (19 spp.), both with as yet numerous undescribed species (MacFarlane et al. 2010; Bickel 2023). Despite this great extant diversity, no fossil representatives of Empididae were previously reported from New Zealand.
Locality and methods
Locality
The specimen described herein was collected at the Foulden Maar Fossil-Lagerstätte, Otago, southern New Zealand (45.5269°S, 170.2191°E). Laminated (varved) and highly fossiliferous diatomite at this locality accumulated in a small (~2000 m diameter) and deep (100–200 m) maar lake over a period of at least 130.000 years in the earliest Miocene (Lindqvist and Lee 2009; Kaulfuss 2017). Anoxic conditions on the profundal lakebed facilitated exquisite preservation of a diverse fossil flora (leaves, flowers and inflorescences, fruits and seeds) and fauna (freshwater sponges, spiders, insects, fish) from a lake/rainforest ecosystem (Bannister et al. 2012; Mildenhall et al. 2014; Kaulfuss et al. 2015; Lee et al. 2016; 2022; Selden and Kaulfuss 2018). The forest growing around the palaeolake, i.e. the palaeohabitat of the Hilarempis specimen described herein, was a Lauraceae-dominated rainforest growing under warm-temperate to subtropical climate, very similar to modern simple notophyll vine forests of eastern Australia (Bannister et al. 2012; Mildenhall et al. 2014). The open lake edge vegetation included some quick-growing tree species in the Mallotus-Macaranga complex on the lakeshore and aquatic plants such as bur reeds, bulrushes and flaxes growing in shallow, swampy areas (Lee et al. 2010; Mildenhall et al. 2014)
Data from 40Ar/39Ar dating, pollen biostratigraphy and sediment magnetic properties indicate that Foulden Maar erupted in the latest Oligocene at ~23.3 Ma and that sedimentation of the ~190 m thick lacustrine crater sequence lasted at least 130.000 years until the earliest Miocene (Lindqvist and Lee 2009, Mildenhall et al. 2014, Fox et al. 2015, Kaulfuss 2017). The Hilarempis specimen was collected in the uppermost 10 m of the preserved diatomite section, which correspond to the latest uppermost Rhoipites waimumuensis to lower early Proteacidites isopogiformis Zone (Aquitanian; New Zealand local stages: late Waitakian–early Otaian). The fossil locality is registered as I43/f8503 in the New Zealand Fossil Record File (https://fred.org.nz/) administered by the Geoscience Society of New Zealand and GNS Science.
Methods
The fossil specimen was studied and photographed with a Nikon SMZ1000 stereomicroscope equipped with a Canon T3 camera. Wetting the fossil with ethanol enhanced the contrast and visibility of anatomical details. Photomicrographs taken at up to 15 focal planes were stacked using Photoshop CS5.1 software (Adobe Systems Inc.). Terms used for adult structures follow those of Cumming and Wood (2017).
Systematic Palaeontology
Order Diptera
Family Empididae Latreille, 1804
Subfamily Empidinae Latreille, 1804
Tribe Hilarini Collin, 1961
Genus Hilarempis Bezzi, 1905
Type species Hilarempis nudifacies Bezzi, 1905
Hilarempis otagoensis sp. nov.
Habitus of Hilarempis otagoensis sp. nov. from Foulden Maar, male, specimen OU47068. Scale bar = 2 mm
Studied material: OU47068, male, Geology Museum, Department of Geology, University of Otago.
Etymology: The specific epithet refers to the Otago region where the specimen was found.
Type locality and horizon: Foulden Maar near Middlemarch, Otago, New Zealand; earliest Miocene (Aquitanian, New Zealand local stages: late Waitakian–early Otaian).
Description: Male. Both wings, left halter, thorax, abdomen with genitalia and one leg (possibly midleg) visible. Thorax apparently darker than abdomen (Fig. 1).
Wing 3.27 mm long, broad with angular anal lobe; Sc evanescent, ending beyond origin of R2+3; apical section of R1 expanded; radial fork broad, with R4 bell-shaped; cell dm shorter than length of M1 (Figs. 2–3).
Hilarempis otagoensis sp. nov. from Foulden Maar, close-up of wing. Scale bar = 1 mm
Hilarempis otagoensis sp. nov. from Foulden Maar, line drawing of wing. Veins: C costa; CuA anterior branch of cubitus; CuP posterior branch of cubitus; M media; R radius; Sc subcosta. Cells: cua anterior cubital; bm basal medial; br basal radial; dm discal medial. Scale bar = 1 mm
Midleg (?) with femur and tibia subequal in length; apical third of tibia with setae, shorter than width of tibia; length of tarsomere 1 subequal to length of tarsomeres 2 and 3 combined; apical tarsomeres missing.
Male terminalia upright, typical Hilarini-type; smaller than width of abdomen, higher than long; phallus tubular.
Remarks: The fossil is assigned to the tribe Hilarini, which is defined in part by the apically thickened (or swollen) vein R1, which is clearly visible in the fossil. Some diagnostic characters for tribal classification and differentiation of genera within Empidinae are wing venation (vein Sc complete/incomplete), male basitarsus I swollen/ unmodified, presence/ absence of laterotergal setae, the setae on tibia I of both sexes, structure of hypopygium and genitalia and morphology of the antennae (Bickel 1996, Grimaldi and Cumming 1999, Sinclair and Cumming 2006). Taken individually many of these characters can be variable in expression and genera (and tribes) of Empidinae are ideally defined by a set of these characters (e.g. Bickel 1996). Although very few diagnostic characters are decipherable in the fossil from Foulden Maar, it readily falls within the rather broad definition of the genus Hilarempis based on the vein Sc being incomplete (Bickel 1996, Sinclair and Cumming 2006). It thus differs from the cosmopolitan genus Hilara, which has a complete vein Sc, with a strongly curved apex.
Discussion
New Zealand has a sparse but increasing fossil record of Diptera. From the Eocene at Livingstone, Otago, Harris (1983) described the march fly Dilophus campbelli Harris (Bibionidae), based on a larva, the only insect found at this site. Other fossil Diptera have been reported from late Oligocene and Early Miocene amber (Schmidt et al. 2018) and the mid-Miocene Hindon Maar Complex (Kaulfuss et al. 2018) in Otago, South Island. Families represented at these sites include Bibionidae, Cecidomyiidae, Ceratopogonidae, Chironomidae, Muscidae(?) and Mycetophilidae but detailed descriptions of individual taxa are not yet available. Diptera known from Foulden Maar include a few adult specimens resembling Tipulidae and Muscidae or Acalyptratae groups (Kaulfuss et al. 2015) and a fauna of pre-imaginal aquatic Chironomidae, Chaoboridae and brachyceran flies (Baranov et al. 2024).
Hilarempis otagoensis sp. nov. described here from Foulden Maar is the first fossil representative of dance flies (Empididae) from New Zealand and the first fossil record of this genus globally (Evenhuis 1994). Extant species of Hilarempis often form dense swarms over water, with males skimming the water surface looking for prey and nuptial gifts. A swarming behaviour can be assumed for males of the fossil Hilarempis species from Foulden Maar, based on its presence in lacustrine sediments of a small, isolated volcanogenic lake. Its more or less articulated preservation suggests that this species lived in the immediate surroundings of the lake, probably among riparian vegetation on the lakeshore. As most extant empidid flies (e.g. MacFarlane et al. 2010), both adults and larvae may have preyed on other insects, mainly other Diptera.
Hilarempis is a widespread genus but likely not monophyletic and it differs from the genus Hilara by a small venational change. Together with the limited diagnostic details preserved in the fossil this prevents any biogeographical comparisons. The new fossil provides unequivocal evidence for the presence of empidid flies in the tribe Hilarini in southern New Zealand in the earliest Miocene, concurring with the great diversity of this genus in the extant New Zealand fauna.
Data availability
This manuscript has no associated data. The material is housed at the Geology Museum, Department of Geology, University of Otago.
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Acknowledgements
We thank the Gibson family at Foulden Hills for kindly allowing access to the fossil site and Daphne Lee for coordinating research at Foulden Maar over many years. We thank Daniel J. Bickel and an anonymous reviewer for critical comments, which improved an earlier version of the manuscript.
Funding
Open Access funding enabled and organized by Projekt DEAL. Palaeontological research at Foulden Maar was financially supported by a Royal Society of New Zealand Marsden Fund (awarded to D.E. Lee, University of Otago). This study was funded by the German Research Foundation (DFG project 429296833).
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Kaulfuss, U., Sinclair, B.J. A dance fly (Empididae: Hilarempis Bezzi) from the Foulden Maar Fossil-Lagerstätte (Early Miocene, New Zealand). Palaeobio Palaeoenv (2024). https://doi.org/10.1007/s12549-024-00606-5
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DOI: https://doi.org/10.1007/s12549-024-00606-5