New Zealand Natural Sciences (2018) 43:1-20 © New Zealand Natural Sciences
Available online at www.science.canterbury.ac.nz/nzns
An annotated checklist of New Zealand mayies
(Ephemeroptera), 2018
Stephen R. Pohe
1, 2
*
1
Pohe Environmental, Whangarei, New Zealand
2
School of Biological Sciences, University of Canterbury, Christchurch, New
Zealand
* Corresponding author: Pohe.Environmental@gmail.com
(Received May 2018, revised and accepted July 2018)
Abstract
e New Zealand mayy (Ephemeroptera) fauna currently comprises 55 described
species belonging to 20 genera in eight families. All species and genera are endemic
to New Zealand, as are three families. e purpose of this paper is to draw together
the fragmented information of published literature in the form of a checklist of
New Zealand Ephemeroptera. e checklist is annotated with species authority and
publication details, type specimen information (sex, life stage, type locality, collector,
specimen repository), distribution and conservation status data, and previously
recognised names, including synonyms. References to all valid species descriptions
and previously recognised names are included in the bibliography.
Keywords: New Zealand, mayies, Ephemeroptera, type localities, type speci-
mens, species distributions, conservation status
Introduction
The mayflies (Ephemeroptera) are an
ancient insect lineage dating back over
300 million years and are believed to
be the most primitive group of extant
winged insects (Grimaldi & Engel 2005;
Bauernfeind & Soldán 2012). ey are
represented on all continents except
Antarctica, but have strong distributional
affinities to specific biogeographical
regions (Gressitt 1967; Barber-James
et al. 2008). Only three families have
global distributions, the remaining 39
being fairly evenly distributed between
the northern and southern hemispheres.
Globally, about 3,600 mayy species have
been described (Barber-James et al. 2013;
M. Sartori and H. Barber-James pers.
comm., May 2018; P. Suter pers. comm.,
May 2018), making the Ephemeroptera
one of the smaller orders of insects.
The New Zealand mayfly fauna
comprises 55 described species in 20
endemic genera and eight families,
three of which are also endemic to New
Zealand (Ichthybotidae, Rallidentidae,
Siphlaenigmatidae). They have close
phylogenetic relationships with the
cold-adapted mayies of southern South
America and Australia, land masses
which like New Zealand, were parts
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New Zealand Natural Sciences 43 (2018)
these authors. A new genus and species
of Leptophlebiidae, Aupouriella pohei, was
also described by Winterbourn (2009).
Other valuable New Zealand mayy con-
tributions include the distribution maps
of specimens held in the Canterbury Mu-
seum (Hitchings 2001; Hitchings et al.
2015) and a chapter in Volume 2 of ‘e
New Zealand Inventory of Biodiversity’
(Gordon 2010), which provides an excel-
lent summary of the New Zealand mayy
fauna and includes a list of 48 species.
e purpose of this paper is to draw
together fragmented information from
the published literature in the form of
a checklist of New Zealand Ephemer-
optera. e checklist is annotated with
species authority and publication details,
type specimen information (sex, life
stage, type locality, collector, specimen
repository), distribution and conservation
status data, and previously recognised
names, including synonyms. References
to all valid species descriptions, and to
previously recognised names, are included
in the bibliography.
Format of the checklist
The higher classification (suborder
and family) of the New Zealand
Ephemeroptera used in this paper is
based on “McCaerty’s system” presented
in Ogden and Whiting (2005, see Figure
2B). However, it is acknowledged that
the status of the suborders Pisciforma
and Setisura are in question (Ogden
et al. 2009) and may be subject to
change. The higher classification is
presented rst, followed by an annotated
checklist of genera and species in each
family. Binomial names are listed in
alphabetical order by family, genus and,
when recognised, subgenus. ese are
annotated with species authority and
publication page number, and with type
of the ancient continent of Gondwana
(Edmunds 1975; Tsui & Peters 1975;
Gibbs 2016). e rst mayies described
from New Zealand were Coloburiscus
humeralis (Walker, 1853) (as Palingenia
humeralis) and Neozephlebia scita (Walker,
1853) (as Baetis scita). In the following 50
years, nine valid species were described
by Eaton (1871, 1883–1888, 1899) and
McLachlan (1873, 1894). Eaton (1899)
also provided the rst annotated check-
list of New Zealand Ephemeroptera (as
Ephemeridae), which included 11 species.
Subsequently, Hudson (1904) published
a popular account of the biology of New
Zealand aquatic insects that also included
one new mayfly species, and Tillyard
(1923) added two more. Phillips (1930)
made a comprehensive revision of the
New Zealand mayy fauna, including
descriptions of eight new species, but in
the following 50 years only four more
species were described (Lestage 1935;
Penniket 1962, 1966; Peters 1971).
Checklists of the aquatic and water-
associated insects of New Zealand, which
included mayflies, were published by
Wise (1965, 1973) who also produced
a synonymic checklist of the smaller
hexapod orders that listed 27 mayfly
species (Wise 1977). Included amongst
a large series of papers on the New Zea-
land Leptophlebiidae, Towns and Peters
(1979a, b) and Towns (1983b) described
eight species in seven genera, and eight
further species were included in their
subsequent monograph (Towns & Peters
1996), which elevated the total number
of New Zealand mayies to 42. More
recently, a revision of the Nesameletidae
by Hitchings and Staniczek (2003) added
three species of Nesameletus. Eight species
of Deleatidium (Hitchings 2008, 2009,
2010; Hitchings & Hitchings 2016) and
a species of Rallidens (Staniczek & Hitch-
ings 2014) have also been described by
3
Pohe. New ZealaNd Mayfly CheCklist 2018
specimen information if known (sex, life
stage, type locality, collector, specimen
repository). Specimen data are followed
by recognised species distributions
and conservation status classications.
Distributions are indicated by the two
letter locality codes of Crosby et al.
(1998) (Figure 1) and North, South, and
Oshore Island localities are separated by
two solidus symbols (//). Conservation
status classications are sourced directly
from the most recent listings for New
Zealand freshwater invertebrates
(Grainger et al. 2014). Finally, other
names by which valid species have been
known are given, and it is stated whether
these names were by original designation
(orig.), synonyms (syn.) or the result of
recombination (comb.). Further, notes
are also given to indicate that a name
was published without a description of
the taxon (undesc.), was misspelt (spell.),
or was the result of misidentification
(misid.). Reference details are provided
for each entry.
Documentation of species in the above
format enables one to search for any
published name, including synonyms,
and it provides easy access to relevant
publication information including page
number, type data, known distributions
and conservation status. Distribution
data provided for each species are based
on records in the literature, records from
ocial New Zealand databases (National
Rivers Water Quality Network) and
collections (New Zealand Arthropod
Collection, Museum of New Zealand,
Canterbury Museum, Otago Museum),
and data from a nation-wide New
Zealand mayfly field survey of adults
and nymphs, undertaken by the author
during 2013–2016. e combined data-
set from all sources currently comprises
73,459 records and forms the basis of an
inaugural national mayy database pres-
ently under construction by the author.
However, it is by no means exhaustive, as
material exists in several national collec-
tions that are yet to be assessed, i.e., part
of the New Zealand Arthropod Collec-
tion, the Auckland Museum Collection
and the NIWA Collection.
Status of type material in Canterbury
Museum, Museum of New Zealand and
New Zealand Arthropod Collection were
conrmed by their respective collection
curators. However, at the time of publica-
tion no response had been received from
e Natural History Museum, London,
regarding type material held there. In
addition, the type repository of Colobur-
iscus tonnoiri Lestage, 1935 is currently
unknown but “may be in a museum in
Brussels” (Terry Hitchings, Canterbury
Museum Research Fellow, pers. comm.,
11 April 2018). e following abbrevia-
tions are used to indicate repositories of
type material listed in the checklist:
BMNH - e Natural History Museum,
London, England
CMNZ - Canterbury Museum,
Christchurch, New Zealand
NMNZ - Museum of New Zealand/
Te Papa Tongarewa, Wellington, New
Zealand
NZAC - New Zealand Arthropod
Collection, Landcare Research, Auckland,
New Zealand
4
New Zealand Natural Sciences 43 (2018)
Figure 1. Map of New Zealand with area codes used to describe specimen distribution records.
For more details, including oshore island area codes, see Crosby et al. (1998).
5
Pohe. New ZealaNd Mayfly CheCklist 2018
Higher classication of New Zealand Ephemeroptera
Class: Insecta
Order: Ephemeroptera Hyatt & Arms, 1890: 69
Suborder: Furcatergalia
Family: Ichthybotidae Demoulin, 1957: 336
Family: Leptophlebiidae Banks, 1900: 246 (as Leptophlebini)
Suborder: Pisciforma
Family: Ameletopsidae Edmunds, 1957: 246 (as Ameletopsinae)
Family: Nesameletidae
1
Kluge et al., 1995: 115
Family: Oniscigastridae Lameere, 1917: 62 (as Oniscigastrina)
Family: Rallidentidae Penniket, 1966: 169 (as Rallidentinae)
Family: Siphlaenigmatidae Penniket, 1962: 394
Suborder: Setisura
Family: Coloburiscidae Edmunds in Edmunds et al., 1963: 11
(as Coloburiscinae)
Species checklist of New Zealand Ephemeroptera
AMELETOPSIDAE
Ameletopsis Phillips, 1930: 324
Ameletopsis perscitus (Eaton, 1899): 291, “♀ imago, No. 26. Wellington (Hudson)”.
Holotype: BMNH. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, RI, WN,
WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD //
SI. Conservation status: Not reatened.
• Ephemera, n.s., near Coloburus (undesc.) (Hudson 1892: 105)
• Ameletus perscitus (orig.) (Eaton 1899: 291)
• Ameletopsis percitus (spell.) (Mosely 1932: 6)
COLOBURISCIDAE
Coloburiscus Eaton, 1888: 332, 346
Coloburiscus humeralis (Walker, 1853): 552, “♀ imago, New Zealand (Sinclair)”.
Holotype: BMNH. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, HB, RI, WI,
WN, WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL,
FD // SI. Conservation status: Not reatened.
• Palingenia humeralis (orig.) (Walker 1853: 552)
• Baetis remota (syn.) (Walker 1853: 564)
• Coloburus humeralis (comb.) (Eaton 1868: 89)
• Coloburus [= Coloburiscus] humeralis (comb.) (Eaton 1888: 332)
1
Also Riek, 1973: 164 (as Nesameletinae)
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New Zealand Natural Sciences 43 (2018)
Coloburiscus tonnoiri Lestage, 1935: 353 (nomen dubium), “♂ imago, Nekotupu
2
,
New Zealand (Tonnoir)”. Holotype: repository unknown. Distribution: AK // – .
Conservation status: Taxonomically Indeterminate & Data Decient.
ICHTHYBOTIDAE
Ichthybotus Eaton, 1899: 285
Ichthybotus bicolor Tillyard, 1923: 228, “♂ imago, Maitai River, Nelson (Philpott)”.
Holotype: Was Cawthron Institute, Nelson, now NZAC. Distribution: – // SD, NN,
BR, WD, MB, KA, NC, MC, SC, OL, CO, DN. Conservation status: Not reatened.
Ichthybotus hudsoni (McLachlan, 1894): 270, “Wellington (Hudson)”. No holotype
designated. Lectotype: “♂ imago, Wellington, (Hudson)”, BMNH. Distribution:
ND, AK, CL, WO, BP, TK, TO, GB, HB, RI, WI, WN, WA // – . Conservation
status: Not reatened.
• Ephemera hudsoni (orig.) (McLachlan 1894: 270)
LEPTOPHLEBIIDAE
Acanthophlebia Towns, 1983: 28
Acanthophlebia cruentata (Hudson, 1904): 33, “Wellington”. No holotype
designated. Lectotype: “♀ subimago, Campbell’s Stream, Karori, Wellington
(Hudson)”, NMNZ. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, HB, RI,
WN, WA // – . Conservation status: Not reatened.
• Atalophlebia cruentata (orig.) (Hudson 1904: 33)
• Zephlebia (Zephlebia) cruentata (comb.) (Penniket 1961: 9)
Arachnocolus Towns & Peters, 1979: 444
Arachnocolus phillipsi Towns & Peters, 1979: 446, “♂ imago, Cascade Stream,
Auckland (Towns)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TK
// – . Conservation status: Not reatened.
• Gen. nov. sp. A (undesc.) (Towns 1976: 46, Table 17; —1978a: 367, 369;
—1978b: 411)
Atalophlebioides
3
Phillips, 1930: 336
Atalophlebioides cromwelli (Phillips, 1930): 385, “Streams round Wellington”.
No holotype designated. Lectotype: “♀ imago, Hutt River, Wellington (inferred as
Phillips)”, NMNZ. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, HB, RI,
WN // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD.
Conservation status: Not reatened.
• Deleatidium (Atalophlebioides) cromwelli (orig.) (Phillips 1930: 385)
2
Likely Nihotupu in the Waitakere Ranges, Auckland (see Craig et al. 2012, page 104).
3
A revision of Atalophlebioides was also published by Towns and Peters (1978).
7
Pohe. New ZealaNd Mayfly CheCklist 2018
Aupouriella Winterbourn, 2009: 424
Aupouriella pohei Winterbourn, 2009: 428, “♂ imago, Whiriwhiri Stream, North
Cape (Pohe)”. Holotype: CMNZ. Distribution: ND // – . Conservation status:
Nationally Critical.
Austroclima Towns & Peters, 1979: 213
Austroclima jollyae Towns & Peters, 1979: 220, “♂ imago, Cascade Stream, Auckland
(Towns)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TK, TO, WN
// SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD // SI.
Conservation status: Not reatened.
• Atalophlebioides sp. B (undesc.) (Towns 1978b: 411; —1979: 257)
Austroclima sepia (Phillips, 1930): 383, “Streams round Wellington”. No holotype
designated. Neotype: “♂ imago, Cascade Stream, Auckland (Towns)”, NZAC.
Distribution: ND, AK, CL, WO, BP, TK, TO, GB, HB, RI, WI, WN, WA // SD, NN,
BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD // SI. Conservation
status: Not reatened.
• Deleatidium (Atalophlebioides) sepia (orig.) (Phillips 1930: 383)
• Atalophlebioides sepia (comb.) (Peters & Edmunds 1964: 238)
Austronella Towns & Peters, 1996: 23
Austronella planulata (Towns, 1983): 17, “♂ imago, Glow-worm Grotto, Waitomo
Caves, Waikato (Pugsley)”. Holotype: NZAC. Distribution: ND, AK, WO, BP, TK,
TO, GB, HB, RI, WN, WA // – . Conservation status: Declining.
• Zephlebia (Zephlebia) planulata (orig.) (Towns 1983b: 17)
Cryophlebia Towns & Peters, 1979: 230
Cryophlebia aucklandensis (Peters, 1971): 47, “♂ imago, rocky stream, Bivouac
Hill, Mt Eden, Auckland Islands (Gressitt)”. Holotype: NZAC. Distribution: AI.
Conservation status: Naturally Uncommon.
• Atalophlebioides aucklandensis (orig.) (Peters 1971: 47)
Deleatidium Eaton, 1899: 288
Deleatidium (Deleatidium) acerbum Hitchings & Hitchings, 2016: 55, “♂ imago,
Camp Creek, Westland (Hitchings & Hitchings)”. Holotype: CMNZ. Distribution:
– // NN, BR, WD. Conservation status: Not yet assessed.
Deleatidium (Deleatidium) angustum Towns & Peters, 1996: 33, “♂ imago, Cascade
Stream, Auckland (Towns)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP,
TK, TO, GB, HB, RI, WN, WA. Also unveried reports from SD, NN, WD, KA,
MC, SC, CO, SL. Conservation status: Not reatened.
• Deleatidium myzobranchia (misid.) (Towns 1979: 255–256)
• Deleatidium nr myzobranchia (undesc.) (Towns 1981: 193–194, 197)
• Deleatidium sp. A (undesc.) (Towns 1983b: 41–47; —1985: 233; —1987:
353–356)
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New Zealand Natural Sciences 43 (2018)
Deleatidium (Deleatidium) atricolor Hitchings, 2009: 40, “♂ imago, Edwards
River, St James Range (Hitchings)”. Holotype: CMNZ. Distribution: – // BR, WD,
MB, KA, NC, MC, SC, MK, OL, CO, SL, FD. Conservation status: Not reatened.
Deleatidium (Deleatidium) autumnale Phillips, 1930: 371, “R. Waikanae and R.
Hutt, Wellington district”. No holotype designated. Lectotype: “♂ imago, Hutt River,
Wellington (Phillips)”, CMNZ. Distribution: ND, WO, BP, TK, TO, GB, HB, RI,
WN, WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL,
FD. Conservation status: Not reatened.
• Deleatidium autumnale (orig.) (Phillips 1930: 371)
Deleatidium (Deleatidium) branchiola Hitchings, 2009: 45, “♂ imago, Mataura
River tributary, Hector Mts (Hitchings)”. Holotype: CMNZ. Distribution: – // OL,
CO. Conservation status: Naturally Uncommon.
Deleatidium (Deleatidium) cerinum Phillips, 1930: 382, “R. Hutt, Wellington
district”. No holotype designated. Lectotype: “♂ imago, Hutt River, Wellington
(Phillips)”, CMNZ. Distribution: ND, AK, WO, BP, TK, TO, GB, RI, WN, WA //
SD, NN, BR, WD, MB, NC, MC, SC, MK, OL, CO, DN, SL, FD // SI. Conservation
status: Not reatened.
• Deleatidium cerinum (orig.) (Phillips 1930: 382)
Deleatidium (Deleatidium) fumosum Phillips, 1930: 372, “Streams round
Wellington”. No holotype designated. Neotype: “♂ imago, Hutt River at Kaitoke,
Wellington (Towns)”, NZAC. Distribution: ND, AK, CL, WO, BP, TK, TO, GB,
RI, WN, WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN,
SL, FD // SI. Conservation status: Not reatened.
• Deleatidium fumosum (orig.) (Phillips 1930: 372)
• Deleatidium sp. C (undesc.) (Towns 1979: 257; —1981: 194; —1983a: 40–47;
—1985: 233; —1987: 353)
Deleatidium (Deleatidium) kawatiri Hitchings & Hitchings, 2016: 61, “♂ imago,
Little Ten Mile Creek, Buller (Hitchings)”. Holotype: CMNZ. Distribution: – // NN,
BR. Conservation status: Not yet assessed.
Deleatidium (Deleatidium) kiwa Hitchings, 2010: 33, “♂ imago, Borland Burn,
South Branch tributary, Fiordland (Ward & Ward)”. Holotype: CMNZ. Distribution:
– // SL, FD. Also unveried report from SI. Conservation status: Naturally Uncommon.
Deleatidium (Deleatidium) lillii Eaton, 1899: 289, “Dunedin (Lillie) and Wellington
(Hudson, No. 46)”. Holotype missing. Lectotype: “♂ imago, No. 46. Wellington
(Hudson)”, BMNH. Distribution: ND, AK, CL, WO, BP, TK, TO, RI, WN, WA
// SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD // SI.
Conservation status: Not reatened.
• Atalophlebia scita (misid. & incorrectly attributed to Walker) (Lillie 1898: 167)
• Deleatidium lillii (orig.) (Eaton 1899: 289)
• Deleatidium (D.) lillii (incorrectly attributed to Walker) (Phillips 1930: 336, 368)
• Deleatidium lilli (spell. & incorrectly attributed to Walker) (Mosely 1932: 10)
• Deleatidium lillii (comb.) (Ulmer 1938: 105)
9
Pohe. New ZealaNd Mayfly CheCklist 2018
Deleatidium (Deleatidium) magnum Towns & Peters, 1996: 40, “♂ imago,
Whakapapaiti Stream, Mt Ruapehu (Crawford)”. Holotype: NZAC. Distribution:
TK, TO. Also unveried reports from NN, NC, MC, SC, MK, CO. Conservation
status: Naturally Uncommon.
Deleatidium (Deleatidium) myzobranchia Phillips, 1930: 373, “Hawkes Bay,
Wellington, Nelson and Canterbury provincial districts”. No holotype designated.
Lectotype: “♂ imago, Ngaio, Wellington (Phillips)”, CMNZ. Distribution: ND, AK,
CL, WO, BP, TK, TO, RI, WN, WA // SD, NN, BR, WD, MB, KA, NC, MC, SC,
MK, OL, CO, DN, SL, FD // SI. Conservation status: Not reatened.
• Deleatidium myzobranchia (orig.) (Phillips 1930: 373)
• Deleatidium sp. E (undesc.) (Towns 1979: 257)
Deleatidium (Deleatidium) townsi Hitchings, 2009: 36, “♂ imago, Glentui River,
Mt omas Forest (Ward & Ward)”. Holotype: CMNZ. Distribution: – // SD, NN,
BR, WD, MB, KA, NC, MC, SC, MK. Conservation status: Not reatened.
Deleatidium (Deleatidium) vernale Phillips, 1930: 360, “Tributary of the Kaiwarra
4
Stream at Ngaio and Khandallah near Wellington”. No holotype designated. Lectotype:
♂ imago, Ngaio, Wellington (Phillips)”, CMNZ. Distribution: ND, CL, TO, GB,
RI, WN, WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN,
SL, FD. Conservation status: Not reatened.
• Deleatidium vernale (orig.) (Phillips 1930: 360)
Deleatidium (Deleatidium) wardorum Hitchings, 2010: 28, “♂ imago, Waterfall
Creek, Ashley (Hitchings)”. Holotype: CMNZ. Distribution: – // SD, NN, BR,
WD, MB, KA, NC, MC, SC, MK, OL, CO. Conservation status: Not reatened.
Deleatidium (Penniketellum
5
) cornutum Towns & Peters, 1996: 47, “♂ imago,
stream at base of Tasman Glacier (Towns & Towns)”. Holotype: NZAC. Distribution:
– // BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, FD. Conservation status:
Not reatened.
Deleatidium (Penniketellum
5
) insolitum (Towns & Peters, 1979): 451, “♂ imago,
Edwards Valley, nr Arthur’s Pass, North Canterbury (Jackson)”. Holotype: NZAC.
Distribution: – // NC, MK. Conservation status: Nationally Endangered.
• Penniketellus insolitus (orig.) (Towns & Peters 1979a: 451)
Deleatidium (Penniketellum
5
) patricki Hitchings, 2008: 37, “♂ imago, Flush
Stream, Mt St. Bathans, Central Otago (Patrick & Edwards)”. Holotype: CMNZ.
Distribution: – // NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, FD.
Conservation status: Not reatened.
4
Kaiwharawhara Stream, Wellington.
5
I here follow the nomenclature of Towns and Peters (1996) but acknowledge the spelling surrounding
the gender may be incorrect. e original designation “Penniketellus” of Towns and Peters (1979a) may
be the valid designation.
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New Zealand Natural Sciences 43 (2018)
Isothraulus Towns & Peters, 1979: 439
Isothraulus abditus Towns & Peters, 1979: 442, “♂ imago, Small tributary of
Waitakere R. nr Anderson’s Track, Auckland (Black)”. Holotype: NZAC. Distribution:
ND, AK, CL, WO, BP, TK, TO, RI // – . Conservation status: Declining.
• Zephlebia sp. A (undesc.) (Towns 1978b): 410
Mauiulus Towns & Peters, 1979: 224
Mauiulus aquilus Towns & Peters, 1996: 51, “♂ imago, Rangitukia Stream, Waikato
(Summerhays)”. Holotype: NZAC. Distribution: ND, CL, WO, TO, WA // – .
Conservation status: Naturally Uncommon.
Mauiulus luma Towns & Peters, 1979: 226, “♂ imago, Cascade Stream, Auckland
(Towns)”. Holotype: NZAC. Distribution: ND, AK, WO, BP, TK, TO, GB, RI, WN
// NN, BR, WD, MB. Conservation status: Not reatened.
• Atalophlebiodes sp. A (undesc.) (Towns 1978a: 367, 369; —1978b: 411; —1979:
256)
Neozephlebia Penniket, 1961: 9
Neozephlebia scita (Walker, 1853): 570, “New Zealand”. No holotype designated.
Lectotype: one of the original two specimens, “♂ imago lacking head, New Zealand
(Sinclair)”, BMNH. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, RI, WN,
WA // SD, NN, BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD //
SI. Conservation status: Not reatened.
• Baetis scita (orig.) (Walker 1853: 570)
• Leptophlebia nodularis (syn.) (Eaton 1871: 81)
• Leptophlebia scita (comb.) (Eaton 1871: 81)
• Atalophlebia nodularis (syn.) (Eaton 1884: 89; —1899: 288; Lillie 1898: 168;
Hutton 1898: 216; Hudson 1904: 34; Phillips 1930: 352; Mosely 1932: 8;
Kimmins 1960: 295; McLean 1967: 99–104)
• Atalophlebia scita (comb.) (Eaton 1884: 90; —1899: 288; Hutton 1898: 216;
Hudson 1904: 34; Kimmins 1960: 295)
• Zephlebia (Neozephlebia) nodularis (syn.) (Penniket 1961: 9; Winterbourn &
Towns 1981: 18)
• Zephlebia (Neozephlebia) scita (comb.) (Penniket 1961: 9; Winterbourn &
Towns 1981: 18)
• Zephlebia scita (comb.) (Cadwallader 1975a: 12, 16, 22; —1975b: 16, 20–21;
—1975c: 304–308; Michaelis 1977: 366)
• Zephlebia scita/nodularis (syn.) (Towns 1978a: 367)
• Zephlebia nodularis (syn.) (Towns 1978b: 410; —1979: 255)
Tepakia Towns & Peters, 1996: 54
Tepakia caligata Towns & Peters, 1996: 55, “♂ imago, Waikoha Stream, Waikato
(Summerhays)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TK, TO,
GB, WN, WA // – . Conservation status: Not reatened.
• ”a second species of Isothraulus” (undesc.) (Towns & Peters 1979a: 444)
11
Pohe. New ZealaNd Mayfly CheCklist 2018
Zephlebia Penniket, 1961: 8
Zephlebia borealis (Phillips, 1930): 356, “Tanekaha, west of Hikurangi, North
Auckland”. No holotype designated. Lectotype: “slides of nymphal legs and gills,
designated from paratype with no locality data (Phillips)”, BMNH. Distribution: ND,
AK, CL, WO, BP, TK, TO, GB, RI, WN // – . Conservation status: Not reatened.
• Atalophlebia ? n. sp. [also provisionally as A. borealis] (orig.) (Phillips 1930: 356)
• Zephlebia (Zephlebia) borealis (comb.) (Penniket 1961: 9)
• Zephlebia (Zephlebia) sp. (cf. borealis) (comb.) (Landa et al. 1980: 171)
• Zephlebia (Terama) borealis (comb.) (Towns 1983b: 19)
Zephlebia dentata (Eaton, 1871): 80, “New Zealand”. No holotype designated.
Lectotype: “♂ imago, designated by Kimmins (1960) with locality as New Zealand
(collector unknown)”, BMNH. Distribution: ND, AK, CL, WO, BP, TK, TO, GB,
RI, WI, WN, WA. Also recorded from the upper South Island but may be ‘in error’.
Conservation status: Not reatened.
• Leptophlebia dentata (orig.) (Eaton 1871: 80)
• Atalophlebia dentata (comb.) (Eaton 1884: 88; —1899: 287; Hutton 1898: 215;
Hudson 1904: 31; Phillips 1930: 344; Mosely 1932: 7; Kimmins 1960: 295)
• Zephlebia (Zephlebia) dentata (comb.) (Penniket 1961: 9)
Zephlebia inconspicua Towns, 1983: 12, “♂ imago, Cascade Stream, Auckland
(Towns)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TO // – .
Conservation status: Not reatened.
• Zephlebia n.sp. B (undesc.) (Towns 1978a: 367)
• Zephlebia sp. B (undesc.) (Towns 1976: 44; —Towns 1978a: 369; —1978b: 410)
• Zephlebia (Zephlebia) inconspicua (orig.) (Towns 1983b: 12)
Zephlebia nebulosa Towns & Peters, 1996: 63, “♂ imago, Rangitukia Stream, Waikato
(Summerhays)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TK, TO // – .
Also unveried reports from BR and SD. Conservation status: Naturally Uncommon.
• Zephlebia sp. A (undesc.) (Towns 1987: 352–353)
Zephlebia pirongia Towns & Peters, 1996: 65, “♂ imago, Rangitukia Stream,
Waikato (Summerhays)”. Holotype: NZAC. Distribution: WO // – . Conservation
status: Not reatened.
Zephlebia spectabilis Towns, 1983: 14, “♂ imago, Waitakere River, Auckland
(Black)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP, TK, TO, GB, RI,
WN, WA // SD, NN, BR, WD, MB, KA, MC, SC, OL, CO, DN, SL, FD // SI.
Conservation status: Not reatened.
• Zephlebia (Zephlebia) sp. (undesc.) (Towns 1979: 256; —1981: 194)
• Zephlebia (Zephlebia) spectabilis (orig.) (Towns 1983b: 14)
Zephlebia tuberculata Towns & Peters, 1996: 68, “♂ imago, Rangitukia Stream,
Waikato (Summerhays)”. Holotype: NZAC. Distribution: ND, AK, CL, WO, BP,
TK, GB, RI // – . Conservation status: Data Decient.
12
New Zealand Natural Sciences 43 (2018)
Zephlebia versicolor (Eaton, 1899): 286, “Wellington (Hudson)”. No holotype
designated. Lectotype: “♂ imago, Wellington (Hudson)”, BMNH. Distribution: ND,
AK, CL, WO, BP, TK, TO, GB, RI, WN, WA // SD, NN, BR, MB. Conservation
status: Not reatened.
• Atalophlebia versicolor (orig.) (Eaton 1899: 286; Hudson 1904: 30; Phillips
1930: 339; Mosely 1932: 7; Kimmins 1960: 296; Stout 1969: 491; —1973:
244; —1975: 442)
• Zephlebia (Zephlebia) versicolor (comb.) (Penniket 1961: 8)
NESAMELETIDAE
Nesameletus Tillyard, 1933: 11
Nesameletus austrinus Hitchings & Staniczek, 2003: 24, “♂ imago, Camp Stream,
Craigieburn Range (Hitchings)”. Holotype: CMNZ (unavailable for conrmation at
time of publication). Distribution: – // NN, BR, WD, MB, KA, NC, MC, SC, MK,
OL, CO, SL, FD. Conservation status: Not reatened.
• Nesameletus sp. A (undesc.) (Winterbourn 2003: 41–50)
Nesameletus avitinctus (Tillyard, 1923): 226, “♀ imago, Waihi Stream, near
Tokaanu (Tillyard)”. Holotype: Was Cawthron Institute, Nelson, now NZAC.
Distribution: ND, AK, CL, WO, BP, TK, TO, GB, HB, RI, WN // SD, NN, BR,
WD, MB, KA, NC, MC, SC, OL, CO, DN. Conservation status: Not reatened.
• Ameletus avitinctus (orig.) (Tillyard 1923: 226)
Nesameletus murihiku Hitchings & Staniczek, 2003: 22, “♂ imago, Mill Creek,
Stewart Island (Hitchings)”. Holotype: CMNZ. Distribution: – // OL, CO, SL, FD
// SI. Conservation status: Naturally Uncommon.
Nesameletus ornatus (Eaton, 1883): plate XIX, “Christchurch (Wakeeld)”. Holotype
missing. Neotype: “♀ imago, Wellington (Hudson, No. 42)”, BMNH. Distribution:
ND, AK, CL, WO, BP, TK, TO, GB, HB, RI, WN, WA // SD, NN, BR, WD, MB,
KA, NC, MC, SC, MK, OL, CO, DN, SL, FD. Conservation status: Not reatened.
• Chirotonetes (?) ornatus (orig.) (Eaton 1883: plate XIX; —1885: 208; —1888:
321)
• Ameletus ornatus (comb.) (Eaton 1899: 291, plate X)
Nesameletus vulcanus Hitchings & Staniczek, 2003: 26, “♂ imago, Narbey
Stream, Banks Peninsula (Hitchings)”. Holotype: CMNZ. Distribution: – // MC.
Conservation status: Nationally Vulnerable.
ONISCIGASTRIDAE
Oniscigaster McLachlan, 1873: 109
Oniscigaster distans Eaton, 1899: 293, “Wainui-o-mata River, Wellington (Hudson,
No. 34 and 34b)”. No holotype designated. Lectotype: “♀ imago, Wainuiomata R.,
N.Z. (Hudson, No. 34)”, BMNH. Distribution: TK, TO, GB, RI, WN // SD, NN,
BR, WD, MB, KA, NC, MC, SC, MK, OL, CO, DN, SL, FD // SI. Conservation
status: Not reatened.
13
Pohe. New ZealaNd Mayfly CheCklist 2018
Oniscigaster intermedius Eaton, 1899: 292 (nomen dubium), “♀ imago, M’Arthur
6,
Nelson, 3600 feet (Hudson, No. 34a)”. Holotype: BMNH. Distribution: – // NN.
Conservation status: Data Decient.
Oniscigaster wakeeldi McLachlan, 1873: 110, “Christchurch, N.Z.”. No holotype
designated. Lectotype: “♀ imago, No. 20. Christchurch, N.Z. (Wakeeld)”, BMNH.
Distribution: ND, WO, TK, TO, GB, WN, WA // SD, NN, BR, WD, MB, KA,
NC, OL, CO, DN, SL. Conservation status: Declining.
RALLIDENTIDAE
Rallidens Penniket, 1966: 164
Rallidens mcfarlanei Penniket, 1966: 164, “♂ imago, Waipoua, North Auckland
(Penniket)”. Holotype: CMNZ. Distribution: ND, AK, CL, WO, BP, TK, TO, RI,
WA // – . Also unveried reports from BR. Conservation status: Not reatened.
Rallidens platydontis Staniczek & Hitchings, 2014: 2, “♂ imago, Otaio River trib.,
e Hunters Hills (Morris)”. Holotype: CMNZ. Distribution: – // NN, MB, NC,
SC, OL, DN, SL. Conservation status: Declining.
SIPHLAENIGMATIDAE
Siphlaenigma Penniket, 1962: 389
Siphlaenigma janae Penniket, 1962: 390, “♂ imago, Baxter’s Creek, Stillwater,
Westland (Penniket)”. Holotype: CMNZ. Distribution: ND, AK, CL, BP, TK, TO,
GB, RI // BR. Conservation status: Nationally Vulnerable.
• Siphlaenigma edmundsi (syn.) (Lugo-Ortiz & McCaerty 1998: 210)
6
Mount Arthur, Nelson.
14
New Zealand Natural Sciences 43 (2018)
Concluding remarks
Fifty-ve New Zealand mayies have been
described, two of which are presently best
considered nomina dubia (Coloburiscus
tonnoiri Lestage, 1935 and Oniscigaster
intermedius Eaton, 1899). The fauna
comprises eight families, including three
endemic to New Zealand (Ichthybotidae,
Rallidentidae, Siphlaenigmatidae), and
four shared with Southern Hemisphere
continents of Gondwanan origin. e
New Zealand species have been described
over a 165 year period and can be
categorised into ve distinct eras based
on the time of their description: pre 1905
(12 species), 1920s & 1930s (11 species),
1960s & 1970s (8 species), 1980s and
1990s (11 species) and post 2000 (13
species). Each era is characterised by the
activity of a small number of passionate
taxonomists who collectively described
0.2–0.7 species per year (Figure 2). In
the most recent period (the last 18 years)
Mr Terry Hitchings, an honorary research
fellow at Canterbury Museum and his
co-authors, have described 13 new species
and been almost solely responsible for
advancing mayfly taxonomy in New
Zealand.
e sporadic nature and low rate of
New Zealand mayy descriptions is not
a reflection of the actual biodiversity
present, but is more likely attributable to
the lack of value (and nancial support)
placed on taxonomic work by New Zea-
land policy makers and biodiversity man-
agers. A lack of support for taxonomic
work is not limited to the Ephemeroptera
by any means and has been expressed
more broadly by New Zealand ento-
mologists (Lester et al. 2014) and aquatic
scientists (Collier et al. 2016), as well as
by taxonomists in other parts of the world
(see Winterbourn 2014). A general ab-
sence of specialist aquatic insect taxono-
mists in New Zealand, and particularly
at New Zealand universities, has meant
that graduates with taxonomic skills are
not being produced, which does not
0
10
20
30
40
50
60
1840 1860 1880 1900 1920 1940 1960 1980 2000
2020
seiceps debircsed fo rebmuN
Date
1.
2.
3. 4. 5.
Figure 2. Cumulative frequency of New Zealand mayy species described over time. Pale blue
bars indicate the ve eras whose primary contributors are: 1. F. Walker, A. Eaton, R. McLachlan & G.
Hudson; 2. R. Tillyard & J. Phillips; 3. J. Penniket & W. Peters; 4. W. Peters & D. Towns; 5. T. Hitchings.
15
Pohe. New ZealaNd Mayfly CheCklist 2018
bode well for the future of fundamental
taxonomic studies of insects, including
aquatic taxa, in this country.
at such studies are needed is well
illustrated by the mayies, as a number of
undescribed species, mainly in the genera
Deleatidium, Nesameletus and Zephlebia
are known to be present in both the
North and South islands, and preliminary
molecular analyses of some described
New Zealand species indicate the possible
presence of cryptic species in these and
other genera (author’s unpublished data).
Taxonomic studies combining morpho-
logical and molecular data are therefore
needed to better understand the diversity
of the New Zealand mayy fauna.
Acknowledgements
I thank Matthew Shaw and Terry
Hitchings (Canterbury Museum), Phil
Sirvid (Museum of New Zealand),
Cody Fraser (Otago Museum), Darren
Ward and Grace Hall (New Zealand
Arthropod Collection) and John Early
(Auckland Museum) for enabling me
to examine mayy collections in their
care, and/or for providing information
on their collection material. Matthew
Shaw, Phil Sirvid and Grace Hall are
also acknowledged for conrming the
status of type material in their respective
collections. I express my appreciation
to Helen Barber-James, Michel Sartori
and Phil Suter for information on
world mayfly diversity. Pieter Pelser
and Michael Winterbourn (University
of Canterbury), Richard Leschen (New
Zealand Arthropod Collection) and Brian
Smith (NIWA Hamilton) are thanked
for helpful comments that improved the
manuscript. I also thank the editors Katie
Collins and Brandon Goeller, and two
anonymous reviewers, for their eorts
and comments.
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