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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3955 (2): 245
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http://dx.doi.org/10.11646/zootaxa.3955.2.4
http://zoobank.org/urn:lsid:zoobank.org:pub:828A05F3-D367-498A-ABDB-70E1126EAE47
Zemacrosaldula, a new genus of Saldidae (Hemiptera: Heteroptera)
from New Zealand: taxonomy, geographic distribution, and biology
MARIE-CLAUDE LARIVIÈRE & ANDR
É
LAROCHELLE
New Zealand Arthropod Collection, Landcare Research, Private Bag 92170, Auckland 1142, New Zealand.
E-mail: LariviereM@landcareresearch.co.nz; LarochelleAndre@hotmail.com
Abstract
Zemacrosaldula, new genus, is described with Salda australis White, 1876, as type species, resulting in the following new
combination Zemacrosaldula australis (White, 1876). Three new species are described: Zemacrosaldula kapekape new
species, Z. whakarunga new species, Z. pangare new species. A revision of the taxonomy of all taxa is presented. Species
are keyed. Morphological descriptions are provided together with illustrations emphasising the most significant diagnostic
features of external morphology and male genitalia. Information is given on synonymy, type specimens, material exam-
ined, geographic distribution and biology.
Key words: shore bugs, revision, systematics, biodiversity
Introduction
Relatively little is known about the shore bugs (Hemiptera: Saldidae) of New Zealand. Seven endemic species so
far placed in the genus Saldula Van Duzee, 1914, have been described from this country (Larivière & Larochelle
2004; 2014). No taxonomic revision or identification key has ever been published for this fauna.
In addition to original species descriptions, the most useful contributions to the taxonomy and general
understanding of the New Zealand Saldidae have been made by Cobben (1961; 1980a), Polhemus (1985a), and
Schuh & Polhemus (2009). Cobben (1961) published descriptions for three species, doubling the number of New
Zealand species assigned to Saldula. Cobben’s descriptions were accompanied by useful illustrations but no
identification key; an informative discussion on the composition of the fauna was however provided. Cobben’s
(1980a) revision of Hawaiian Saldula was also insightful especially in terms of dealing with “taxonomic
complications” caused by character polymorphism (e.g., variation in hemelytral pigmentation or in degree of
development of hindwings) which, as he put it, can hardly be resolved using single specimens or the small samples
found in most museums. Polhemus (1985a) published the most comprehensive world overview of the family
Saldidae, including a preliminary generic-level phylogeny. Consequently his work yielded the most exhaustive
framework to consider regional faunas. Without the broad perspective offered by Polhemus (1985a) it would have
been extremely difficult to undertake this New Zealand study mainly because the Southern Hemisphere fauna in
general and the Australian fauna in particular, remain largely unresolved taxonomically. Schuh & Polhemus’s
(2009) revision and phylogenetic analysis of the South American and Andean genus Pseudosaldula contributed
further useful contextual information for the New Zealand revision, especially with respect to morphological
character analysis, saldid phylogeny, and austral biogeography.
The Australian Saldidae (three genera, ten species) are only partially known taxonomically and the most
relevant literature on the subject (Rimes 1951; Cobben 1980b; Polhemus 1991; Cassis & Gross 1995) so far does
not suggest a particularly close affinity with the New Zealand fauna.
The proposition that the New Zealand saldids described to date are not congeneric with Saldula, is not new
(Polhemus 1985a–b; Lindskog & Polhemus 1992; Schuh & Polhemus 2009) but little could be done until now to
revise this classification for lack of sufficient and well-documented material. Extensive fieldwork by the authors
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since 1992 has yielded large amount of new saldid material, including long series of specimens from several
populations and a wealth of new information on geographic distribution and biology. This provided solid bases
upon which to conduct the current study.
The genus Saldula (sensu lato) is a large, nearly worldwide, polyphyletic taxon that has often served as a
‘wastebasket genus’ to classify species that could not readily fit elsewhere. Lindskog & Polhemus (1992) estimated
that nearly half of known Saldidae were contained in Saldula and that nearly half of Saldula species could
eventually be transferred to other genera. These authors were the first to suggest a more restricted taxonomic
definition and description of Saldula (sensu stricto), using a monophyletic concept based on two synapomorphies
(median cell of corium with eye spot; median endosomal sclerite of male with anteromedial laminar projections).
The taxa revised here do not fit Lindskog & Polhemus’s definition of Saldula or the taxonomic concept of
other described saldid genera. Consequently a new genus, Zemacrosaldula (Saldidae: Saldinae: Saldoidini), is
erected to accommodate Salda australis White, 1876 (formerly placed in Saldula ) as well as three new species (Z.
kapekape, Z. whakarunga, Z. pangare). This brings the total number of known saldid taxa for New Zealand to two
genera, Zemacrosaldula and Saldula (sensu lato), containing four and six species respectively.
Zemacrosaldula species are easily recognised in the field. They are the largest of the New Zealand Saldidae.
Their overall body colour is blackish with variously shaped pale dorsal hemelytral markings and speckles. They are
generally heliophilous, showing peak activity during intense sunlight periods, and mostly saxicolous, living on
stones, gravel and other rocky formations arising from or located near freshwater. They live along riverbanks—
including the long braided rivers of the South Island—and on the shores of lakes. This is typically a group of inland
species that can at times be found in estuaries near the mouths of mainly large rivers as long as suitable riparian
habitat is available and upstream populations are well established. Zemacrosaldula species are macropterous, with
well-developed hindwings, and able to fly at least short distances.
The authors hope that their efforts to clarify the alpha-taxonomy of species in this genus and to publish
extensive biological and distributional information can address some of the limitations to advancing knowledge on
New Zealand saldids as well as provide a foundation for more detailed systematics and evolutionary studies.
Materials and methods
This study is based on the examination of over 1,100 specimens (mostly adults) from more than 130 New Zealand
localities. Most of this material (c. 85%) was collected by the authors from 1992 to 2014 and is deposited in the
New Zealand Arthropod Collection (NZAC), Auckland.
Other specimens were kindly provided by the following New Zealand institutions: Auckland Institute and
Museum, Auckland (AMNZ); BMNH (The Natural History Museum, London, U.K.; formerly British Museum of
Natural History); Canterbury Museum, Christchurch—P.M. Johns specimens (CMNZ); Entomology Research
Museum, Lincoln University, Lincoln (LUNZ); Museum of New Zealand Te Papa Tongarewa, Wellington
(MONZ).
The NZAC specimens used in this study received unique barcode labels and were databased. Once this paper is
published, specimen records will be made available through Landcare Research’s Systematics Collections
Database portal (http://scd.landcareresearch.co.nz/ ).
Terms particular to Saldidae morphology mostly follow Schuh & Polhemus (2009), the most notable
exceptions are as follows: preocellar furrow(s), “sillons antéocellaires” of Péricart (1990); preocular spot(s),
adopted here to denote the rather smooth area of cuticle near the point of insertion of each of a pair of
trichobothrium-like setae near the preocellar furrow; acetabulum (acetabula) of Cobben (1961), the area of the
sternum forming each thoracic cavity into which a leg is inserted, numbered I, II and III (pro-, meso-, metathoracic
acetabulum); inner membrane of parandria, adopted here instead of medial membrane. For the male aedeagus, the
expression anterolateral sclerites refers to the apicolateral sclerites of Polhemus (1985a).
The adjective pruinose is used to qualify dull hemelytral surfaces that appear ‘frosted’ or ‘covered by a
powdery layer.’ Schuh & Polhemus (2009) showed these pruinose or dull surfaces to bear dense mats of
microtrichia and non-pruinose or shiny surfaces to be devoid of them.
The male genitalia of representatives of as many populations as possible were dissected in the following
manner. Dry-mounted specimens were warmed for 15 minutes in hot soapy water; softened specimens were
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transferred to a watch glass containing some of the hot water used for softening (e.g., square 4 cm watch glass or
equivalent). For each specimen, the genital capsule was separated from the abdomen using fine forceps and a
microscalpel (e.g., needle tip from a 1.0 ml disposable hypodermic syringe or other fine needle); parameres and
aedeagus were detached and extracted from the genital capsule using the same tools; dissected genitalia were
transferred to smaller individual watch glasses (e.g., 1 dram glass vials cut in half) containing glycerol and a thin
film of 75% ethanol to assist transfer of structures from water to glycerol; the small watch glasses were then stored
in 24 wells sorting trays for further examination. At the end of the study dissected genitalia were transferred to
plastic microvials containing glycerol, and subsequently mounted on the pin below relevant specimens.
Descriptions are based on adults. Measurements included in the descriptions, were taken as follows: body
length, in dorsal view, from visible apex of head to apex of hemelytron; antennal segment length, from base to apex
of segment; antenna length, the sum of individual segment lengths; leg segment length, from base to apex of
segment; pronotum or scutellum length, along midline, from base to apex. Cells in the membrane of the hemelytron
are numbered from 1 to 4, from most anterior cell (near costal margin) to most posterior cell (near apex of clavus).
The eunomy or eunomic series—the range of variation in hemelytral pigmentation arranged in a sequence
from light to dark according to a more or less stable gradation pattern for a given species—is an important
taxonomic character used in saldid taxonomy. The eunomic series in this revision represent the general pattern that
best fits variations observed among populations of a species.
Type data, when provided, are listed in this order: type status followed by sex, acronym of entomological
collection or museum serving as repository, and original label data with a forward slash (/) separating data from
different labels.
The taxonomic arrangement of species and illustrations follows the order of taxa in the identification key.
Photographs were taken using Leica stereomicroscopes (MZ16 or M205A), digital cameras (Leica DFC450 or
Nikon DSRi1), and the image stacking applications Helicon Focus or Zerene Stacker. Further photo-processing
was done using Adobe Photoshop, CorelDRAW graphics suite, and Adobe Lightroom. Black and white
illustrations were prepared and laid out using CorelDRAW graphics suite. The distribution map was digitally traced
from a draft map generated from georeferenced locality data using internet-based GIS technology.
The two-letter abbreviation codes of Crosby et al. (1976; 1998) for areas of New Zealand, were used to record
localities. North Island: AK, Auckland; BP, Bay of Plenty; CL, Coromandel; GB, Gisborne; HB, Hawke’s Bay;
ND, Northland; RI, Rangitikei; TK, Taranaki; TO, Taupo; WA, Wairarapa; WI, Wanganui; WN, Wellington; WO,
Waikato. South Island: BR, Buller; CO, Central Otago; DN, Dunedin; FD, Fiordland; KA, Kaikoura; MB,
Marlborough; MC, Mid Canterbury; MK, Mackenzie; NC, North Canterbury; NN, Nelson; OL, Otago Lakes; SC,
South Canterbury; SD, Marlborough Sounds; SL, Southland; WD, Westland. Stewart Island: SI. For each species
areas codes are listed from north to south and west to east. Table 1 provides geographical coordinates of collecting
localities in decimal degrees.
Biological notes are based on an analysis and synthesis of specimen label data and field observations by the
authors.
Taxonomy, geographic distribution, and biology
Zemacrosaldula new genus
Typ e s pecie s . Salda australis White, 1876, by present designation.
Description. Body length 4.22–6.28 (5.15) mm; elongate-ovate or broad-ovate. Dorsal colour somewhat speckled;
largely blackish, with uniformly dark or, more rarely, narrowly pale lateral margins of pronotum, and a few to
several, coalesced or individual pale markings on hemelytra, including a more or less defined line of three to four
spots along R vein. Macropterous. Head not closely appressed to thorax (eyes distinctly separated from thorax),
with three pairs of long trichobothrium-like setae (two dorsally on mandibular plates, two near preocellar furrows,
two near preocellar spots). Frons with well-developed longitudinal furrow medially. Ocelli slightly elevated from
surface of vertex, separated by the diameter of one ocellus or less, closer to each other than to eyes. Preocellar spots
distinct, usually paler than surrounding area, subtriangular to crescent-shaped, narrowly to broadly touching eyes
(by one ocellus width or more), slightly extending or not in front of ocelli. Preocellar furrows present, about as
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deep as longitudinal frontal furrow. Preocular spots indistinct, of same colour as surrounding area. Transverse
swelling (postclypeus of Cobben) rather flat, slightly to moderately developed; lateral portions separated by a gap
(mostly) or contiguous near facial midline. Mandibular plates rather flat, evanescent to moderately developed (not
prominent or tumid). Maxillary plates slightly to moderately developed. Antennae largely dark; segment II
completely dark or narrowly pale subapically only, at least 2.2x longer than I, clothed over entire length with very
short setae (shorter than segment width), without long setae in apical half; segments III–IV not wider than apex of
segment II, with short setae and a number of longer setae (subequal to segment width). Thorax. Pronotum: 0.51–
0.62x as long as scutellum medially; subtrapezoidal; lateral margins subrectilinear to slightly convex, more rarely
sinuate-concave, uniformly dark or, more rarely, narrowly pale (pale area narrower than or about as wide as
antennal segment II), not explanate and not separated from disc by a longitudinal furrow; collar present, continuous
(not interrupted medially), delimited posteriorly by a row of punctures; calli not strongly raised although well
differentiated from disc, contiguous, forming a transverse elevation with a median pit, delimited posteriorly by a
row of punctures curving forward laterally and reaching lateral margins of pronotum. Underside: xyphus 2 elongate
(as long as or longer than wide), subtriangular. Legs: hind tibiae with subapical comb; hind tarsal segment II 1.0–
1.4x as long as segment III. Hemelytra: without a distinct eyespot subbasally on exocorium; pubescence short,
mostly semi-erect, whitish, more or less evenly distributed; clavus and corium largely pruinose or, more rarely,
with pruinosity reduced on corium; costal margin slightly convex along entire length; costal fracture slightly
curved forward, nearly reaching apex of R vein; embolar modification of female moderately (Z. pangare) to
strongly developed; subapical pale spot of clavus present or absent; membrane with four well-formed cells, cell 2
about as long as cell 3, cell 4 distinctly longer than cell 3; hypocostal ridge simple; secondary hypocostal ridge
absent. Abdomen. Venter largely black (never narrowly or broadly pale laterally) with or without posterior margin
of segments narrowly pale. Paired eversible glands present; when externally visible, one gland located on each side
near posterior margin of segment VII. Male parandria (Figs 19–20) elongate, narrowly or broadly subtriangular;
tip acuminate or obtusely rounded; inner membrane present along basal quarter to basal third of inner margins.
Male paramere (Figs 15–18) with or without distinct processus sensualis; dorsal margin bearing moderate to long
setae. Male aedeagus containing three pairs of sclerites (Fig. 23, Z. australis), two larger median branched sclerites
and 4 smaller simple anterolateral sclerites. Male filum gonopori coiled approximately 1.25 or 1.5 times (Figs 24–
25). Female subgenital plate (segment VII, ventrally) with posterior margin produced caudally, truncate medially.
Ovipositor with gonapophyse 1 distinctly serrate. Spermathecal pump flange present. Gynandrial gland sclerotised.
Remarks. The generic name Zemacrosaldula is derived from Ze- (New Zealand) and Macrosaldula (Greek,
makros = large; Latin, Saldula, feminine gender); the name of the Northern Hemisphere genus that Zemacrosaldula
resembles. The authors think that this is the genus tentatively identified as Genus A by Polhemus (1985a), and said
to represent the closest morphological and ecological analogue of Macrosaldula Southwood & Leston, 1959.
The main morphological characters unifying the species of Zemacrosaldula are listed here, with generic level
apomorphies from Polhemus (1985a) in italics: size relatively large (4.22–6.28 mm); dorsal colour somewhat
speckled, largely blackish, with uniformly black or narrowly pale lateral margins of pronotum, and a few to several
coalesced or individual pale markings on hemelytra, including a line of three to four spots along R vein; hemelytra
lacking a distinct eyespot subbasally on exocorium; antennal segment II long, clothed over entire length with very
short setae, without long setae in apical half; frons with well-developed longitudinal furrow medially; calli of
pronotum with a distinct median pit and delimited posteriorly by a row of punctures reaching lateral margins;
hemelytra and hindwings fully developed; membrane with four well-formed cells; embolar modification of female
hemelytra well developed; hind tibiae with subapical comb; male aedeagus with anterolateral sclerites; processus
sensualis of male paramere distinct or indistinct; filum gonopori of male coiled 1.25 or 1.5 times.
The male abdominal grasping plate was not studied in detail but preliminary observations in Z. australis
indicate that it bears about 24 spines in the following configuration: an outer row of roughly 12 moderately long
spines and an inner group of roughly 12 longer spines.
Zemacrosaldula species exhibit a high level of eunomic variability. Divergence from the ‘standard’ eunomy is
common even within populations, and although it is possible to ‘perceive’ the identity of a species based on
hemelytral pigmentation patterns, further confirmation is required from other morphological characters, including
male genitalia. By the same token study material used for character analysis must consist of long series of
specimens from any given population so as to sufficiently cover the range of morphological variations.
Cobben (1980a), in is study of Hawaiian Saldula, noted that environmental factors exert an influence on the
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proportion of colour morphs at the population level, e.g., high temperatures and/or lower humidity shift the
eunomic series towards the light side of the sequence while cooler temperatures and/or higher humidity generally
produce darker morphs. The authors have noted similar trends for Zemacrosaldula species, each occupying a
distribution range encompassing broad and complex climatic conditions mainly caused by extensive topographical
variation (e.g., from coastal lowlands to high mountains), wide-ranging temperatures (e.g., warm or cool temperate
to severely cold) and precipitations (e.g., annual rainfalls of 600–4000 mm, and snow in the mountains). The
authors have also observed that the pigmentation of newly emerged adults and females in general, tends towards
the pale side of pigmentation spectrum.
Key to species of Zemacrosaldula
Remark. Additionally helpful but not necessarily exclusive characters are provided between square brackets.
1 Body elongate-ovate (Figs 1–2). Dorsum mostly dull in appearance, with rather evenly distributed, short whitish
pubescence. Lateral margins of pronotum uniformly dark or narrowly pale. Hemelytra with several individual or
coalesced whitish markings (Figs 1–2, 11–12). Face pubescent (Figs 5–8). Male genitalia: parandria narrowly
subtriangular, acuminate at tip, with evenly curved inner margins (Fig. 19); paramere without distinct processus
sensualis (Figs 15–16); apical half of aedeagus with subelongate and bent anterolateral sclerite (Fig. 21); filum
gonopori coiled 1.5 times (Fig. 24). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Body broad-ovate (stocky) (Figs 3–4). Dorsum shinier in appearance (at least head and pronotum contrastingly
shiny), with sparser, less evenly distributed or reduced, short to very short whitish pubescence. Lateral margins of
pronotum uniformly dark. Hemelytra with few individual (not coalesced) whitish markings (Figs 3–4, 13–14).
Face pubescent (Fig. 9) or glabrous (Fig. 10). Male genitalia: parandria broadly subtriangular, obtusely rounded at
tip, with angular inner margins (Fig. 20); paramere with distinct processus sensualis (Figs 17–18); apical half of
aedeagus with Y-shaped anterolateral sclerite (Fig. 22); filum gonopori coiled 1.25 times (Fig. 25) . . . . . . . . . . . 3
2 Antennae: segment II at least 2.4x longer than segment I. Pronotum: lateral margins uniformly dark or narrowly
pale. Hemelytra with several, often coalesced, whitish markings (Figs 1, 11). Male paramere: processus hamatus
acuminate and slightly upturned at tip (Fig. 15). [Distribution: North Island and northeastern South Island.] . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. australis (White)
- Antennae: segment II at most 2.2x longer than segment I. Pronotum: lateral margins uniformly dark. Hemelytra
with several individual (not coalesced) whitish markings (Figs 2, 12). Male paramere: processus hamatus nar-
rowly rounded and subrectilinear at tip (Fig. 16). [Distribution: southernmost areas and west coast of the South
Island.]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. kapekape new species
3 Face pubescent (Fig. 9); slightly to moderately contrasted in colour (with partially pale transverse swelling, max-
illary plates, anteclypeus). Dorsum (Fig. 3) moderately shiny (especially head, pronotum, scutellum), with short,
somewhat sparse and unevenly distributed whitish pubescence. Hemelytra with clavus and corium largely pruin-
ose. Underside of thorax mostly pubescent. Male paramere (Fig. 17): processus sensualis well developed; proces-
sus hamatus narrowly rounded at tip. [Hemelytra (Fig. 13): corium frequently with a pair of semilunate marks in
basal third and a subquadrate medial mark at about midlength. Distribution: South Island, excluding southernmost
areas and the west coast.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. whakarunga new species
- Face glabrous (Fig. 10); not contrasted in colour (entirely black or at most with transverse swelling slightly paler
near sides or facial midline). Dorsum (Fig. 4) very shiny overall, with very short, very sparse (reduced) and quite
unevenly distributed whitish pubescence. Hemelytra with clavus largely pruinose; corium partially pruinose
(inner endocorium, subbasal exocorium). Underside of thorax mostly glabrous. Male paramere (Fig. 18): proces-
sus sensualis slightly developed; processus hamatus broadly rounded at tip. [Hemelytra (Fig. 14): corium largely
immaculate, frequently with a subquadrate medial mark subapically. Distribution: South Island west coast, in the
vicinity of the Fox and Franz Josef glaciers.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. pangare new species
Zemacrosaldula australis (White, 1876) new combination
Salda australis White, 1876: 106. Holotype male (BMNH; seen by Larivière & Larochelle 2004); New Zealand.
Acanthia australis: Kirkaldy, 1909: 27.
Saldula australis: Drake & Hoberlandt, 1950: 7.
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Description. Body length 4.22–5.89 (4.91) mm; elongate-ovate (Fig. 1). Dorsal colour largely black, with
uniformly dark or narrowly pale lateral margins of pronotum and several whitish markings on hemelytra;
frequently with irregular coalesced marks on exocorium. Facial colour (Figs 5–7) slightly to moderately contrasted.
Head, pronotum and scutellum slightly shiny, contrasting slightly against mostly dull hemelytra. Dorsal
pubescence short, reclined to semi-erect, whitish, more or less evenly distributed. Dorsal microsculpture rugulose
on head, pronotum and scutellum. Head black. Face (Figs 5–7) pubescent. Transverse swelling whitish yellow to
pale brown; lateral portions separated by a narrow to rather wide dark gap or, rarely, almost contiguous (nearly
touching) near facial midline. Mandibular plates whitish yellow to pale brown or nearly black. Maxillary plates,
anteclypeus and labrum whitish yellow to pale brown; anteclypeus often marked with brown basally. Antennae
4.07–4.33x longer than pronotum + collar medially, largely dark; segment I whitish yellow to yellowish brown,
with ventral and dorsal sides dark brown to black (often striped), sometimes nearly entirely black; segment II dark
brown to black, often paler in apical fourth, 2.40–2.77x longer than segment I; segments III–IV dark brown to
black, sometimes slightly darker than segment II. Thorax. Lateral margins of pronotum subrectilinear to very
slightly sinuate-concave, uniformly dark or, more rarely, narrowly pale (pale area usually narrower than,
sometimes about as wide as antennal segment II). Underside black, with sharply contrasting broadly or narrowly
pale acetabula I–III (acetabulum III rarely completely dark) and partially (posterior quarter to half) to almost
completely pale lateral margins; pubescence rather dense, silvery, appressed (except for glabrous lateral margins).
Legs moderately dark; fore, mid femora, and often hind femora striped with dark brown to nearly black on ventral
and dorsal sides, along part or entire length, otherwise whitish yellow with some brown spots, or, nearly completely
black; tibiae with dark brown to black dorsal stripe over most of length (mid and hind tibiae often infuscate only)
and black subapical annulus, otherwise whitish yellow with dark spines; hind tibiae 2.70–2.76x longer than tarsal
segments II+III combined; tarsal segment II darkened or not apically, otherwise whitish yellow, segment III brown
in apical half (more rarely completely brown); pretarsal claws long, about half the length of tarsal segment III.
Scutellum black, 1.76–1.91x longer than pronotum + collar medially. Hemelytra: corium (Fig. 11) largely black
with several irregular, often coalesced, whitish markings; endocorium with a line of four pale spots along R vein, a
few additional spots subapically along membrane, and sometimes a longitudinal mark about middle; exocorium
frequently with rather large irregular coalesced markings on basal third to three-quarters and an irregularly shaped
medial mark subapically; colour pattern in female consistent with that of male but often paler overall with broader
markings; pale markings in darker individuals not coalesced, reduced in size and numbers, with line of four spots
still visible but reduced to diminutive spots or speckles; corium in darkest individuals almost entirely black with
only a few speckles; corium and clavus largely pruinose; basal pale spot of clavus present or absent; subapical pale
spot of clavus present or, more rarely, absent; membrane dark brown to black basally with a pale mark near tip of
clavus, brown medially within cells, pale elsewhere, and with dark brown to black veins. Abdomen. Venter black,
with or without posterior margin of most segments narrowly pale (generally more broadly pale in female).
Pubescence rather dense, silvery, appressed in both sexes. Male parandria (Fig. 19) narrowly subtriangular,
acuminate at tip; inner margins evenly curved; basal margin slightly sinuate. Male paramere (Fig. 15) without
distinct processus sensualis; instead, with flat cuticular surface bearing less than 10 setae; processus hamatus
acuminate and slightly upturned at tip. Apical half of male aedeagus (Fig. 21), in lateral view, with 3 main visible
sclerites (elongate median sclerite; subelongate, bent anterolateral sclerite; small, often subtriangular anterolateral
sclerite). Male aedeagal sclerites, in ventral view, as in Fig. 23. Male filum gonopori (Fig. 24) coiled 1.5 times.
Female subgenital plate (segment VII, ventrally) black with apical half pale or plate largely pale. Other characters
as in generic description.
Geographic distribution (Fig. 26). North Island and northeastern South Island.
Material examined. A total of 621 specimens including holotype, from the following localities. North Island.
BP—Kaimai Range, Dickey Flat Road end (NZAC); Papatea, Waiti Stream (NZAC); Urewera National Park,
Waimana River Valley, Pohatu Track, 3.2 km E Tauwhare (NZAC); Waioeka Gorge, N of Oponae and S of Opotiki
(AMNZ). CL—Coromandel Range, Stony Bay (NZAC); Coromandel Range, West of Kirikiri Saddle (NZAC);
Coromandel Range, Tapu River Valley (AMNZ). GB—Karakatuwhero River Valley, by upper ford (NZAC);
Mangatuna (NZAC); Morere (AMNZ); Te Awha Stream, Rangitukia Road (NZAC); Urewera National Park,
Aniwaniwa (NZAC); Urewera National Park, Te Taita a Makaro (NZAC). HB—Kaweka Forest Park, Lawrence
Road access, Tutaekuri River (NZAC); Tangoio (near) (AMNZ); Tukituki River, Pourerere Road, 10km E of
Waipawa (NZAC); Wakarara, upper Waipawa River (NZAC). RI—Kawhatau River, Junction of Toetoe & Potaka
Roads (NZAC); Mangaweka, Rangitikei River bridge (AMNZ); Vinegar Hill, Junction of Vinegar Hill Road &
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Rangitikei River (NZAC). TK—Inglewood, Manganui River (AMNZ); Mangorei (AMNZ); Mount Egmont (N
side) (AMNZ). TO—Lake Taupo (AMNZ); Mount Ruapehu (Whakapapanui Stream above Chateau (AMNZ);
Whakapapanui Stream, dam (AMNZ)); Owhango, Whakapapa River (AMNZ); Taupo (NZAC); Whakapapanui
Stream, Junction of Mahuia Rapid & Highway 47 (NZAC). WA —Dannevirke, Manawatu River, Aerodrome Road
end (NZAC); Makaretu River, 2 km N of Takapau (NZAC); Mangatewainui River, Junction with Gundry Road
(NZAC). WI—Whangaehu River, 5km N of Mangamahu (NZAC). WN—Kimberley Scenic Reserve (S of Levin),
Ohau River (NZAC); Ngatiawa River (NZAC); Otaki (6 km S (NZAC); 6 km SE (NZAC)); Otaki Forks (AMNZ).
WO—Mangaotaki River, S of PioPio (AMNZ). South Island. BR—Buller River, Gowan River Bridge (NZAC);
Maruia River, Maruia Falls (NZAC); Rotokohu, Inangahua River (NZAC); Inangahua area, White Cliffs, Buller
River (AMNZ). KA—Clarence River mouth (NZAC); Clarence River, 12 km inland via Clarence Valley Road
(NZAC); Oaro (LUNZ), Omihi Stream mouth (LUNZ). MB—Pelorus River, Daltons Bridge (NZAC); Wairau
River (2.5 km N of Leatham Road (NZAC); Church Lane end (NZAC)). MC—Highway 73, 15 km W of
Springfield (NZAC); Rakaia River (NZAC); Waimakariri Gorge (CMNZ), N of Waddington (NZAC); Waimakariri
River (CMNZ). NC—Hurunui Bridge (N side) (CMNZ); Junction of Hope River & Highway 7 (NZAC); Kairaki
Beach (CMNZ).
Biology. Altitudinal range. Lowland (mostly) to low montane, more rarely high montane (e.g., central North
Island volcanic plateau). Not usually coastal; may be present in predominantly freshwater habitats located near the
mouth of rivers that also harbour inland populations. Habitat. Occurs along stony-gravelly (or shingled)
riverbanks, riverbeds or, less frequently, lakeshores; saxicolous, found mainly on gravel, stones or boulders near
the water’s edge, usually within 0.5 to 3 m from the waterline, as well as on stones or boulders emerging from the
water; mostly in sandy environments, also in slightly silty or clayish situations; primarily near moderate to fast
running water, also near calmer water (e.g., side river channels or temporary backwaters). Sometimes found on
bare ground patches between stones or boulders near the waterline or, occasionally, near rocky seepages away from
a main river channel. Nymphs live in the same habitat, on the ground surface between and under stones or among
gravel. Seasonality. Adults collected from October to April; mating pairs observed in December; newly emerged
adults (tenerals) collected mostly in December and March; nymphs found from October to March—suggesting
overwintering in the egg stage, nymphal development in the spring and emergence of a new summer generation
from December, possibly with a one or two month delay in southern parts of the distribution range or at higher
altitudes. Food. Predator or scavenger. Behaviour. Jumps from stone to stone, flies short distances (usually less
than 2 m), dashes quickly into the space between stones or into water when disturbed. Heliophilous; hides under
and between stones, in cloudy or rainy weather.
Remarks. Zemacrosaldula australis is the most widely distributed species of this genus. It is the only
Zemacrosaldula species known from the North Island and it occurs also in northern areas of the South Island.
Zemacrosaldula australis closely resembles the South Island endemic Z. kapekape and can be difficult to
distinguish from it although the two species are largely allopatric. In addition to characters of the male genitalia Z.
australis bears the following distinguishing features: hemelytra usually with larger, more coalesced pale markings;
antennae with longer segment II; paramere with more acutely tipped processus hamatus. Dissection of the male
genitalia is necessary to diagnose the two species with certainty, especially when dealing with darker specimens
with reduced hemelytral markings or populations that occur in relatively close proximity (e.g., on the South Island
west coast).
On the North Island Z. australis should not be confused with Saldula trivialis Cobben, 1961 or Saldula
maculipennis Cobben, 1961. These two taxa may be conspecific or part of an unresolved species-complex. They
superficially resemble Z. australis albeit in a diminutive version with highly distinctive, acutely tipped male
parandria (see Fig. 30 in Cobben 1961) and broadly pale sides of the female abdominal venter.
Morphologically speaking, Z. australis is the most highly variable species in this genus. As a general rule this
species is more darkly coloured and somewhat smaller in body size in southern parts of its range, in more
mountainous or generally colder habitats (e.g., central North Island volcanic plateau) although Z. australis is not
typically monticolous. Anomalous looking specimens of Z. australis, somewhat reminiscent of Z. whakarunga,
were observed from several South Island populations, especially in the lowlands and along the coast in a Kaikoura
(KA), North Canterbury (NC), and Mid Canterbury (MC). The identity of these specimens could only be confirmed
when male specimens were available for genitalic dissection.
Variations in facial colour, pubescence, and degree of development of transverse swelling (Figs 5–7)
encountered in this species are similarly observed in Z. kapekape and Z. whakarunga.
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FIGURES 1–4. Dorsal views of Zemacrosaldula species (legs and antennae omitted). Scale bar = 1 mm. (1) Z. australis, (2) Z.
kapekape, (3) Z. whakarunga, (4) Z. pangare.
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FIGURES 5–10. Facial views of Zemacrosaldula species. (5–7) Z. australis, (8) Z. kapekape, (9) Z. whakarunga, (10) Z.
pangare.
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FIGURES 11–12. Eunomy (left corium), with more frequently observed pigmentation patterns indicated by an asterisk (*).
(11) Zemacrosaldula australis (cl = clavus, en = endocorium, ex = exocorium), (12) Z. kapekape.
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FIGURES 13–14. Eunomy (left corium), with more frequently observed pigmentation patterns indicated by an asterisk (*).
(13) Zemacrosaldula whakarunga, (14) Z. pangare.
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FIGURES 15–24. Schematic views of male genitalia. (15–18) Paramere, ventral view. (15) Zemacrosaldula australis, (16) Z.
kapekape (ph = processus hamatus), (17) Z. whakarunga (ps = processus sensualis),
(18) Z. pangare. (19–20) Parandria,
posterior view. (19) Z. australis, (20) Z. whakarunga. (21–22) Anterior half of aedeagus, lateral view. (21) Z. australis, (22) Z.
whakarunga. (23) Sclerites of aedeagus, Z. australis, ventral view. (24–25) Filum gonopori, lateral view. (24) Z. australis, (25)
Z. whakarunga.
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Zemacrosaldula kapekape new species
Zemacrosaldula kapekape Larivière & Larochelle, new species. Holotype: male (NZAC) labelled “NEW ZEALAND OL Mt
[=Mount] Aspiring NP [=National Park], Cameron Flat 300m 4429S 16848E 17.III.2006 Larivière, Larochelle / Stony-
gravelly streambank: bare silty soil; on stones near water / HOLOTYPE [male symbol] Zemacrosaldula kapekape
Larivière & Larochelle, 2015 (red label). Paratypes: 7 males (1 CMNZ, 1 LUNZ, 1 MONZ, 4 NZAC) and 8 females (1
CMNZ, 1 LUNZ, 1 MONZ, 5 NZAC) with same data as holotype, bearing blue paratype labels.
Description. Body length 4.51–5.53 (5.04) mm; elongate-ovate (Fig. 2). Dorsal colour largely black, with
uniformly dark lateral margins of pronotum and several individual (not coalesced) whitish markings on hemelytra;
frequently with a combination of a pair of semilunate or irregularly shaped marks in basal third of corium and
moderate to large irregularly shaped medial mark at about midlength on exocorium. Facial colour (Fig. 8) slightly
to moderately contrasted. Head, pronotum and scutellum moderately shiny, contrasting moderately against mostly
dull hemelytra. Dorsal pubescence short, reclined to semi-erect, whitish, sometimes more sparse than in Z.
australis but still rather evenly distributed. Dorsal microsculpture rugulose on head, pronotum and scutellum.
Head black. Face (Fig. 8) pubescent. Transverse swelling whitish yellow to yellowish brown; lateral portions
separated by a narrow to rather wide dark gap or, sometimes, almost contiguous (nearly touching) near facial
midline. Mandibular plates whitish yellow to yellowish brown. Maxillary plates whitish yellow. Anteclypeus and
labrum whitish yellow to yellowish brown; anteclypeus often marked with brown basally and dark brown to black
along margins; labrum often margined with dark brown to black. Antennae 4.07–4.33x longer than pronotum +
collar medially, largely dark; segment I whitish yellow to yellowish brown, with ventral and dorsal sides dark
brown to black (usually striped); segment II dark brown to black, often shortly paler subapically, 2.16–2.17x longer
than segment I; segments III–IV dark brown to black. Thorax. Lateral margins of pronotum subrectilinear to very
slightly convex, uniformly dark. Underside black, with sharply contrasting broadly pale acetabula I–II (acetabulum
II sometimes narrowly pale), acetabulum III narrowly to very narrowly pale (rarely completely dark), and partially
pale (posterior half) to almost completely pale lateral margins; pubescence rather dense, silvery, appressed (except
for glabrous lateral margins). Legs moderately dark; fore femora striped with dark brown to nearly black on ventral
and dorsal sides, along part or entire length; mid and hind femora thickly marked with brown on ventral and lateral
sides, otherwise whitish yellow with some brown spots, or, nearly completely black; tibiae with dark brown to
black dorsal stripe over most of length and black subapical annulus, otherwise whitish yellow with dark spines;
hind tibiae 2.71–2.87x longer than tarsal segments II+III combined; tarsal segment II slightly darkened or not
apically, otherwise yellowish brown, segment III completely dark brown to black (often paler in basal half of hind
tarsus); pretarsal claws rather short, about a third of the length of tarsal segment III. Scutellum black, 1.82–1.96x
longer than pronotum + collar medially. Hemelytra: corium (Fig. 12) largely black with several irregular,
individual (not coalesced), whitish markings; endocorium with a line of four pale spots along R vein, a few
additional spots subapically along membrane, sometimes with a spot about middle (rarely an irregular longitudinal
mark); exocorium without coalesced irregular markings on basal third to three-quarters (as opposed to Z. australis),
with a number of individual markings, a moderate to large irregularly shaped medial mark at about midlength, and
usually a smaller irregular medial mark subapically; colour pattern in female consistent with that of male but often
paler overall with broader markings; pale markings in darker individuals reduced in size but hardly in numbers,
with line of four visible spots not significantly reduced in size; corium in darkest individuals never almost entirely
black with a few speckles; corium and clavus largely pruinose; basal pale spot of clavus present or absent;
subapical pale spot of clavus present; membrane dark brown to black basally with a pale mark near tip of clavus,
brown medially within cells and along margin, pale within cells on each side of brown medial mark, and with dark
brown to black veins. Abdomen. Venter black, with posterior margin of some or most segments narrowly pale
(generally more broadly pale in female). Pubescence rather dense, silvery, appressed in both sexes. Male parandria
as in Z. australis (see Fig. 19). Male paramere (Fig. 16) without distinct processus sensualis; instead, with flat
cuticular surface bearing less than 10 setae; processus hamatus narrowly rounded and subrectilinear at tip. Apical
half of male aedeagus, in lateral view, with 3 main visible sclerites as in Z. australis (see Fig. 21). Male filum
gonopori coiled 1.5 times as in Z. australis (see Fig. 24). Female subgenital plate (segment VII, ventrally) black
with apical half pale or plate largely pale. Other characters as in generic description.
Geographic distribution (Fig. 26). Southernmost areas and west coast of the South Island.
Material examined. A total of 223 specimens including types, from the following localities. South Island.
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CO—Clutha River (LUNZ); Clutha River, junction with Highway 8A, 9 km SE of Luggate (NZAC); Danseys Pass
(N of) (CMNZ); Nevis Crossing (LUNZ). DN—Otekaieke, Otekaieke River (NZAC). FD—Hunter Mountains,
South Borland River (NZAC). MK—Lake Pukaki, Whale Stream (NZAC). OL—Glenorchy, Buckler Burn
(NZAC); Lake Hawea (AMNZ); Mount Aspiring National Park, Cameron Flat (NZAC). SC—Hakataramea,
Hakataramea River (NZAC); Aparima River (14 km S of Mossburn (NZAC); 4.2 km S of Wreys Bush (NZAC));
Dipton West, Oreti River (NZAC); Winton, Oreti River (NZAC). WD—Harihari, Wanganui River mouth
(CMNZ); Junction of Wanganui River & Highway 6 (NZAC); Lake Mapourika (MONZ); Lake Moeraki (NZAC);
Omoeroa River, S of Franz Josef (AMNZ); Totara River, near Moorhouse Road (NZAC).
Biology. Altitudinal range. Lowland (mostly) to lower montane. Not usually coastal; may be present in
predominantly freshwater habitats located near the mouth of rivers that also harbour inland populations. Habitat.
Occurs along stony-gravelly (or shingled) riverbanks, riverbeds or, less frequently, lakeshores; saxicolous, found
mainly on gravel, stones or boulders near the water’s edge, usually within 0.5 to 3 m from the waterline, as well as
on stones or boulders emerging from the water; mostly in sandy environments, also in slightly silty or clayish
situations; primarily near moderate to fast running water, also near calmer water (e.g., side river channels or
temporary backwaters). Sometimes found on bare ground patches between stones or boulders near the waterline.
Nymphs live in the same habitat, on the ground surface between and under stones or among gravel. Seasonality.
Adults collected from October to March; mating pairs observed in March; newly emerged adults (tenerals) and
nymphs found in March. Food. Predator or scavenger. Behaviour. Jumps from stone to stone, flies short distances
(usually less than 2 m), dashes quickly into the space between stones or into water when disturbed. Heliophilous;
hides under and between stones, in cloudy or rainy weather.
Remarks. The species name is from the Maori word ‘kapekape’, meaning west-south-west. It refers to the
South Island west coast and southernmost distribution of this species and to it being the west-south-west
counterpart of Z. australis.
The external morphology and male genitalia of Z. kapekape closely resemble those of Z. australis from which
it can be difficult to distinguish. The eunomy falls somewhere between that of Z. whakarunga and darker forms of
Z. australis. The configuration of the male paramere is also reminiscent of the condition observed in Z. australis
but differs mainly in the shape of the processus hamatus which is more bluntly rounded and straight at the tip in Z.
kapekape. In addition, this species usually lacks the coalesced hemelytral markings seen in the basal third to three-
quarters of the exocorium in Z. australis and it has a shorter second segment of the antenna. See also Remarks
under Z. australis.
Zemacrosaldula whakarunga new species
Zemacrosaldula whakarunga Larivière & Larochelle, new species. Holotype: male (NZAC) labelled “NEW ZEALAND SD
Carluke, Rai River 50m 4112S 17335E 27.II.2012 Larivière, Larochelle / Sandy-silty gravelly riverbank; within 2m from
water[line] / HOLOTYPE [male symbol] Zemacrosaldula whakarunga Larivière & Larochelle, 2015 (red label).
Paratypes: 6 males (1 LUNZ, 1 CMNZ, 1 MONZ, 3 NZAC) and 5 females (1 LUNZ, 1 MONZ, 3 NZAC) with same data
as holotype, bearing blue paratype labels.
Description. Body length 4.31–6.28 (5.57) mm; broad-ovate (Fig. 3). Dorsal colour largely black, sometimes with
a very faint maroon hue, with uniformly dark lateral margins of pronotum and a few individual (not coalesced)
whitish markings on hemelytra; frequently with combination of a pair of semilunate marks in basal third of corium
and a small to medium size subquadrate medial mark at about midlength on exocorium. Facial colour (Fig. 9)
slightly to moderately contrasted. Head, pronotum, and scutellum strongly shiny, usually contrasting strongly
against mostly dull hemelytra. Dorsal pubescence short, reclined to semi-erect, whitish, somewhat sparse and
unevenly distributed. Dorsal microsculpture slightly rugulose on head, pronotum and scutellum. Head black. Face
(Fig. 9) pubescent. Transverse swelling yellowish brown to almost entirely black; lateral portions contiguous or,
sometimes, separated by a narrow to rather wide dark gap near facial midline. Mandibular plates black or, less
often, yellowish brown. Maxillary plates whitish yellow to yellowish brown, rarely obscure. Anteclypeus and
labrum yellowish brown to dark brown, narrowly to broadly margined with black. Antennae 4.24–4.85x longer
than pronotum + collar medially, largely dark; segment I mostly black with partial or complete yellowish stripe on
one side, sometimes entirely black; segment II black, often shortly paler subapically, 2.18–2.35x longer than
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segment I; segments III–IV black. Thorax. Lateral margins of pronotum subrectilinear to slightly convex,
uniformly black. Underside black, with moderately contrasting broadly or narrowly pale acetabulum I, broadly to
very narrowly pale acetabulum II, narrowly pale to completely dark acetabulum III, and partially (posterior quarter
to half) to completely dark lateral margins; rarely, if ever, with lateral margins almost completely pale; pubescence
moderately dense, silvery, appressed (except for glabrous lateral margins). Legs largely dark; fore, mid, and often
hind femora almost entirely black or thickly striped with black on ventral and often also dorsal side over most of
length, otherwise whitish yellow to yellowish brown with some brown spots; tibiae with dark brown to black dorsal
stripe over most of length (hind tibiae often infuscate only) and black subapical annulus, otherwise whitish yellow
to yellowish brown with dark spines; hind tibiae about 2.77–2.86x longer than tarsal segments II+III combined;
tarsal segment II darkened apically, otherwise whitish yellow to yellowish brown, segment III completely dark
brown to black (often paler in basal half on hind tarsus); pretarsal claws rather short, about a third of the length of
tarsal segment III. Scutellum black, 1.78–1.83x longer than pronotum + collar medially. Hemelytra: costal margin
(Fig. 3) more convex and embolium generally more explanate basally than in Z. australis or Z. kapekape; corium
(Fig. 13) largely black with few irregular, individual (not coalesced), whitish markings; frequently with a pair of
semilunate marks in basal third, one on each side of R vein (as opposed to Z. pangare); endocorium with a line of
four pale spots (often reduced to two or three) along R vein, a few additional speckles (often reduced) subapically
along membrane; exocorium without coalesced markings, with few individual markings, a small to moderately
large subquadrate medial mark at about midlength, and often a smaller irregular medial mark subapically (often
reduced or absent); colour pattern in female consistent with that of male but regularly with larger subquadrate
medial mark at about midlength on exocorium; pale markings in darker individuals strongly reduced in size and
numbers, with line of two to four spots still visible but reduced to speckles; corium in darkest individuals almost
entirely black and barely speckled; corium and clavus largely pruinose; basal and subapical pale spots of clavus
present or absent; membrane brown to black basally, rarely with a pale mark near tip of clavus, brown medially
within cells and around margin, pale within cells on each side of brown medial mark, and with dark brown to black
veins. Abdomen. Venter black, with (both sexes) or without (male) posterior margin of most segments narrowly
pale. Pubescence moderately dense (although sparser than in Z. australis or Z. kapekape), silvery, appressed in both
sexes. Male parandria (Fig. 20) broadly subtriangular, obtusely rounded at tip; inner margins angular; basal
margin straight. Male paramere (Fig. 17) with distinct, well-developed processus sensualis bearing less than 10
setae; processus hamatus narrowly rounded and often slightly upturned at tip. Apical half of male aedeagus (Fig.
22), in lateral view, with 3 main visible sclerites (elongate median sclerite; shorter Y-shaped anterolateral sclerite;
smaller subtriangular anterolateral sclerite). Male filum gonopori (Fig. 25) coiled 1.25 times. Female subgenital
(segment VII, ventrally) plate black with apical quarter to half pale or, sometimes, plate at most narrowly pale
apically. Other characters as in generic description.
Geographic distribution (Fig. 26). South Island, excluding southernmost areas and the west coast.
Material examined. A total of 228 specimens including types, from the following localities. South Island.
BR—Boyle River, Dans Creek Bridge, Engineers Camp (LUNZ); Fox River (Buller County) (LUNZ); Nelson
Lakes National Park (Lake Rotoiti, head (LUNZ); Mount Robert (LUNZ)); Punakaiki, Bullock Creek (CMNZ).
CO—Lowburn Valley mouth (NZAC); The Remarkables, Rastus Burn (LUNZ); Duntroon, Maerewhenua River
(NZAC); Junction of Danseys Pass Road & S Branch of Maerewhenua River (NZAC); Palmerston, Shag River
(NZAC). KA—Hundalee, Conway River (NZAC); Junction of Kowhai River & Postmans Road (NZAC);
Kahutara River mouth (NZAC); Oaro Beach (LUNZ). MC—Banks Peninsula (Birdlings Flat (CMNZ); near Lake
Ellesmere (LUNZ)); Cameron River (LUNZ); Craigieburn Forest Park, Dracophyllum Track (NZAC); Craigieburn
Ski Field (LUNZ); Mount Algidus (NZAC); Porters Pass, Lake Lyndon (CMNZ, NZAC); South Highway 72, W of
Staveley (NZAC); Thomas River, N branch (LUNZ). NC—Arthurs Pass National Park, Poulter River (NZAC);
Lees Valley, Ashley River bridge (LUNZ); Napenape Scenic Reserve, Hurunui River mouth (NZAC); Okuku
River, 5km W of Loburn (NZAC). NN –Aniseed Valley (NZAC); Iron Hill, Iron Lake (LUNZ); Karamea River
mouth (LUNZ); Lake Sylvester (LUNZ); Lake Sylvester, Bushline Hut (LUNZ); Little Sylvester Lake (LUNZ);
Oparara River (NZAC); Pariwhakaoho River mouth (NZAC); Pearce River Resurgence (LUNZ); Pelorus Bridge
Scenic Reserve (NZAC). OL—Dart Hut (NZAC); Greenstone River (NZAC); Headlong Peak (NZAC), South
Basin Stream (NZAC); Peak 2.5 km W of Mount Tyndall (NZAC). SC –Temuka, Opihi River near Waiapi
(CMNZ). SD—Carluke, Rai River (NZAC); Opouri River, junction with Tunakino Valley Road (NZAC). WD—
Mount Aspiring National Park (Arawata Bivouac (LUNZ); Arawata River (LUNZ)); Nelson Creek domain, beside
Nelson Creek (LUNZ).
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Biology. Altitudinal range. Montane (mostly) to subalpine, more rarely lowland. Exceptionally coastal, in
predominantly freshwater habitats located near the mouth of rivers that also harbour inland populations. Habitat.
Occurs along stony-gravelly (or shingled) riverbanks riverbeds or, less frequently, lakeshores; mostly saxicolous;
found mainly on gravel, stones or boulders near the water’s edge, usually within 0.5 to 3 m from the waterline, as
well as on stones or boulders emerging from the water; mostly in sandy environments, also in moderately silty or
clayish situations; primarily near moderate to fast running water, also near calmer water (e.g., side river channels or
temporary backwaters); at higher altitudes, on wet screes and snowmelt patches. Often taken near water, on bare
ground patches between stones or boulders, on bare ground with few scattered stones (e.g., silty, clayish or rather
muddy flats and banks of otherwise stony-gravelly rivers and lakes), or, sparsely vegetated ground (e.g., at the
margin of subalpine lakes). Occasionally collected near rocky seepages away from a main river channel. Nymphs
live in the same habitat, on the ground surface between and under stones or among gravel, or, hiding at the base of
plants. Seasonality. Adults collected from November to March; mating pairs observed in March; newly emerged
adults (tenerals) collected in March; nymphs found from January to March. Food. Predator or scavenger.
Behaviour. Jumps from stone to stone, flies short distances (usually less than 2 m), dashes quickly into the space
between stones or into water when disturbed. Heliophilous; hides under and between stones or at the base of plants,
in cloudy or rainy weather.
Remarks. The species name ‘whakarunga’ is from the Maori and means south or southern. It refers to the
geographic range of this species which is restricted to New Zealand’s South Island where it is widely distributed.
Zemacrosaldula whakarunga is morphologically closer to Z. pangare than to other species in this genus. In
addition to structural differences in the male genitalia, Z. whakarunga has an overall less shiny, more pubescent,
more pruinose and more blackish appearance than Z. pangare. The hemelytral colour pattern, especially the
frequent occurrence of a pair of semilunate marks in the basal third of the corium and a subquadrate medial mark at
about midlength on the exocorium, is most often diagnostic for Z. whakarunga. See also Remarks under Z.
pangare.
In eastern areas of the South Island, Z. whakarunga can be especially difficult to differentiate from darker,
anomalous looking specimens of Z. australis (see Remarks under that species). In such cases as well as for darker
forms of both species, with strongly reduced hemelytral markings, examination of the male genitalia is the only
reliable identification method. Although found in lowland areas, especially along and near the mouth of large rivers
originating from the Southern Alps (e.g., large South Island braided rivers), Z. whakarunga is habitually
monticolous.
Zemacrosaldula pangare new species
Zemacrosaldula pangare Larivière & Larochelle, new species. Holotype: male (NZAC) labelled “NEW ZEALAND WD Jct
Waikukupa Riv[er] & Hwy [=Highway] 6 200m 4326S 17004E 11.III.2007 Larivière, Larochelle / Sandy-silty stream in
glacial moraine entering main river: on water-logged sand with dead leaves & algae / HOLOTYPE [male symbol]
Zemacrosaldula pangare Larivière & Larochelle, 2015 (red label). Paratypes: 13 males (1 CMNZ, 1 LUNZ, 1 MONZ, 10
NZAC) and 10 females (1 CMNZ, 1 LUNZ, 1 MONZ, 7 NZAC) with same data as holotype, bearing blue paratype labels.
Description. Body length 4.65–5.63 (5.10) mm; broad-ovate (Fig. 4). Dorsal colour largely dark brown to black,
with distinctive maroon hue, uniformly dark lateral margins of pronotum, and very few individual (not coalesced)
whitish markings on hemelytra; exocorium nearly immaculate but frequently with a rather large (more rarely
small) subquadrate medial mark subapically. Facial colour (Fig. 10) not contrasted. Head, pronotum and scutellum
strongly shiny, contrasting moderately against mostly shiny hemelytra. Dorsal pubescence very short, reclined to
semi-erect, whitish to slightly golden, very sparse (reduced) and quite unevenly distributed (more so than in Z.
whakarunga). Dorsal microsculpture very slightly rugulose on head; similarly developed although partly lacking
on pronotum and scutellum. Macropterous. Head black. Face (Fig. 10) glabrous (pubescent in other species).
Transverse swelling black or, very rarely, more lightly coloured at sides or near facial midline (but still mostly
black); lateral portions contiguous or, rarely, separated by a narrow to rather wide dark gap near facial midline.
Mandibular plates, maxillary plates, anteclypeus, and labrum black; maxillary plates seldom marked with brown
speckles; anteclypeus and labrum sometimes lighter brown or narrowly marked with brown apically. Antennae
3.98–4.00x longer than pronotum + collar medially, largely dark; segment I black or with apical fourth to half dark
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yellowish brown or pale brown (not distinctly striped as in Z. whakarunga); segment II dark brown to black, often
shortly paler subapically, 2.59–2.61x longer than segment I; segments III–IV dark brown to black. Thorax. Lateral
margins of pronotum subrectilinear to slightly convex, uniformly black. Underside black, with barely contrasting
narrowly to very narrowly pale (rarely completely dark) acetabulum I, narrowly pale to completely dark
acetabulum II, completely dark (rarely very narrowly pale) acetabulum III, and completely dark lateral margins;
rarely, if ever, with lateral margins narrowly pale brown near posterior edge; pubescence very sparse or evanescent
(overall mostly glabrous). Legs largely dark; fore, mid, and usually hind femora entirely black, at most shortly
tinged with dark yellowish brown or pale brown subapically; tibiae with dark brown to black dorsal stripe over
most of length (hind tibiae often infuscate only) and black subapical annulus, otherwise dark yellowish brown with
dark spines; hind tibiae about 2.87–2.91x longer than tarsal segments II+III combined; tarsal segment II not of
barely darkened apically, otherwise yellowish brown, segment III completely dark brown to black (often paler in
basal half on hind tarsus); pretarsal claws rather short, about a third of the length of tarsal segment III. Scutellum
black, 1.61–1.89x longer than pronotum + collar medially. Hemelytra: costal margin (Fig. 4) convex and embolium
basally explanate as in Z. whakarunga; embolar modification of female moderately developed (strongly developed
in other species); corium (Fig. 14) largely black with very few irregular, rather small, individual (not coalesced)
whitish markings; markings lacking in basal third on each side of R vein (as opposed to Z. whakarunga);
endocorium with a line of three pale spots (often reduced to one or two) along R vein, with or without additional
speckles subapically along membrane (as opposed to Z. whakarunga); exocorium without coalesced markings (as
opposed to Z. australis) or a subquadrate mark at about midlength (as opposed to Z. whakarunga and Z. kapekape),
with very few individual markings and a rather large subquadrate medial marking subapically (often reduced,
sometimes absent); colour pattern in female consistent with that of male but regularly with a larger subapical
medial mark on exocorium; corium in darkest individuals almost entirely black and barely speckled; clavus largely
pruinose; corium partially pruinose (inner endocorium, subbasal exocorium, especially embolium); basal pale spot
of clavus absent; subapical pale spot of clavus present or, more rarely, absent; membrane brown to black basally,
without pale mark near tip of clavus, brown medially within cells and around margin, pale within cells on each side
of brown medial mark, and with dark brown to black veins. Abdomen. Venter black, with or without posterior
margin of at least some segments narrowly pale. Pubescence much shorter than in other Zemacrosaldula species,
very sparse along middle, nearly lacking elsewhere, silvery, appressed in both sexes. Male parandria as in Z.
whakarunga (see Fig. 20). Male paramere (Fig. 18) with distinct, slightly developed processus sensualis bearing
less than 10 setae; processus hamatus broadly rounded at tip. Apical half of male aedeagus, in lateral view, with 3
main visible sclerites as in Z. whakarunga (see Fig. 22). Male filum gonopori coiled 1.25 times as in Z.
whakarunga (see Fig. 25). Female subgenital plate (segment VII, ventrally) completely black or at most with apical
half of sides pale. Other characters as in generic description.
Geographic distribution (Fig. 26). South Island west coast, in the vicinity of the Fox and Franz Josef glaciers.
Material examined. A total of 44 specimens including types (NZAC), from the following localities. South
Island. WD—Junction of Clearwater River & Gillespies Beach Road (NZAC); Junction of Waikukupa River &
Highway 6 (NZAC); Franz Josef (NZAC).
Biology. Altitudinal range. Lowland (50–200 m). Habitat. Found along a sandy silty stream in a glacial
moraine entering a main river, on waterlogged sand and seepages with sparse dead leaves and algae (several
specimens, Junction Waikukupa River and Highway 6); in glacial moraine, among cushion plants (one specimen,
Franz Josef); on bare silty ground patch along an otherwise stony riverbank (one specimen, Junction Clearwater
River and Gillespies Beach Road). Apparently not obligatory saxicolous. Seasonality. Adults collected in March;
newly emerged adults (tenerals) also found in March. Food. Predatory or scavenger. Behaviour. Sluggish, jumping
for a distance of 15 cm or less, if jumping at all, when disturbed (several specimens, Junction Waikukupa River and
Highway 6). Presumably heliophilous like other Zemacrosaldula species.
Remarks. The species name ‘pangare’ means beardless in Maori. It refers to the smooth, hairless face of this
species and its overall shinier, more glabrous appearance, compared to other Zemacrosaldula species.
It is mostly the above mentioned appearance and structural differences in the male genitalia that set Z. pangare
apart from Z. whakarunga. The maroon hue of the dorsum, best seen in oblique lateral view, appears mostly due to
the pruinosity of the hemelytra. A similar colour hue can, to some extent, also be seen on the eyes and antennae.
The following characters are also diagnostic for Z. pangare: face almost entirely black; hemelytra lacking markings
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in the basal third of the corium on each side of R vein and bearing a subquadrate medial mark subapically on the
exocorium.
While all other Zemacrosaldula species are primarily saxicolous and live close to the waterline of main river
channels, Z. pangare favours waterlogged soil and ground seepages located at some distance from stony-gravelly
riverbanks and riverbeds. See also Remarks under Z. whakarunga.
FIGURE 26. Collecting localities of Zemacrosaldula species, New Zealand.
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TABLE 1 . Geographical coordinates of localities in decimal degrees.
Locality (standard) AreaCode Latitude Longitude
Aniseed Valley NN -41.3878 173.1608
Aparima River, 14 km S of Mossburn SL -45.7833 168.1833
Aparima River, 4.2 km S of Wreys Bush SL -46.0000 168.1333
Arthurs Pass National Park, Poulter River NC -42.8753 171.8611
Banks Peninsula, Birdlings Flat MC -43.8261 172.7078
Banks Peninsula, near Lake Ellesmere MC -43.7300 172.4400
Boyle River, Dans Creek Bridge, Engineers Camp BR -42.5647 172.3558
Buller River, Gowan River Bridge BR -41.7000 172.5500
Cameron River MC -43.4236 171.0847
Carluke, Rai River SD -41.2000 173.5833
Clarence River mouth KA -42.1667 173.9167
Clarence River, 12 km inland via Clarence Valley Road KA -42.1000 173.8333
Clutha River CO -45.6500 169.4000
Clutha River, junction with Highway 8A, 9 km SE of Luggate CO -44.7500 169.3333
Coromandel Range, Stony Bay CL -36.5167 175.4167
Coromandel Range, West of Kirikiri Saddle CL -37.1628 175.6136
Coromandel Range,Tapu River Valley CL -36.9807 175.5279
Craigieburn Forest Park, Craigieburn Ski Field MC -43.1175 171.7039
Craigieburn Forest Park, Dracophyllum Track MC -43.1594 171.7033
Dannevirke, Manawatu River, Aerodrome Road end WA -40.2333 176.0833
Danseys Pass CO -44.9478 170.3819
Dart Hut OL -44.5219 168.5544
Dipton West, Oreti River SL -45.9000 168.3833
Duntroon, Maerewhenua River DN -44.8500 170.6833
Fox River BR -42.0401 171.4080
Franz Josef WD -43.3887 170.1824
Glenorchy, Buckler Burn OL -44.8500 169.4000
Greenstone River OL -44.9053 168.1617
Hakataramea, Hakataramea River SC -44.7167 170.4833
Harihari, Wanganui River mouth WD -43.1411 170.5547
Headlong Peak OL -44.5403 168.5969
Headlong Peak, South Basin Stream OL -44.5581 168.6078
Highway 73, 15 km W of Springfield MC -43.3167 171.8000
Hundalee, Conway River KA -42.5833 173.4167
Hunter Mountains, South Borland River FD -45.7456 167.3803
Hurunui Bridge (N side) NC -42.8969 173.0978
Inangahua area, White Cliffs, Buller River BR -41.8000 171.8700
Inglewood, Manganui River TK -39.1581 174.2755
Iron Hill, Iron Lake NN -41.1056 172.6155
Junction Hope River & Highway 7 NC -42.5833 172.4500
Junction of Clearwater River & Gillespies Beach Road WD -43.4500 169.9000
Junction of Danseys Pass Road & S Branch of Maerewhenua River DN -44.9333 170.5833
Junction of Kowhai River & Postmans Road KA -42.3667 173.5833
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TABLE 1. (Continued)
Locality (standard) AreaCode Latitude Longitude
Junction of Waikukupa River & Highway 6 WD -43.4333 170.0667
Junction of Wanganui River & Highway 6 WD -43.1500 170.6167
Kahutara River mouth KA -42.4333 173.5833
Kaimai Range, Dickey Flat Road end BP -37.4387 175.7473
Kairaki Beach NC -43.3856 172.7114
Karakatuwhero River Valley, by upper ford GB -37.6761 178.2706
Karamea River mouth NN -41.2561 172.1047
Kaweka Forest Park, Lawrence Road access,Tutaekuri River HB -39.3667 176.3667
Kawhatau River, Junction Toetoe & Potaka Roads RI -39.7833 175.8833
Kimberley Scenic Reserve (S of Levin), Ohau River WN -40.6669 175.3133
Lake Hawea OL -44.6119 169.2615
Lake Mapourika WD -43.3167 170.2164
Lake Moeraki WD -43.7167 169.2667
Lake Pukaki, Whale Stream MK -43.9500 170.1167
Lake Sylvester NN -41.1065 172.6286
Lake Sylvester, bushline Hut NN -41.1060 172.6447
Lake Taupo TO -38.6900 176.0800
Lees Valley, Ashley River bridge NC -43.1856 172.1503
Little Sylvester Lake NN -41.1060 172.6447
Lowburn Valley mouth CO -45.0000 169.2167
Makaretu River, 2 km N of Takapau WA -40.0000 176.3500
Mangaotaki River, S of PioPio WO -38.5209 174.9063
Mangatewainui River, Junction with Gundry Road WA -40.0667 176.1667
Mangatuna GB -38.3019 178.2686
Mangaweka, Rangitikei River bridge RI -39.8247 175.7845
Mangorei TK -39.0893 174.1024
Maruia River, Maruia Falls BR -41.8600 172.2528
Morere GB -38.9809 177.7903
Mount Algidus MC -43.2386 172.2031
Mount Aspiring National Park, Arawata Bivouac WD -44.4136 168.6081
Mount Aspiring National Park, Arawata River WD -44.4064 168.5931
Mount Aspiring National Park, Cameron Flat OL -44.4833 168.8000
Mount Egmont (N side) TK -39.2600 174.0500
Mount Ruapehu, Whakapapanui Stream above Chateau TO -39.2097 175.5446
Mt Ruapehu, Whakapapanui Stream, dam TO -39.1914 175.5312
Napenape Scenic Reserve, Hurunui River mouth NC -42.9139 173.2744
Nelson Creek domain, beside Nelson Creek WD -42.5036 171.6036
Nelson Lakes National Park, Lake Rotoiti, head BR -41.8169 172.8372
Nelson Lakes National Park, Mount Robert BR -41.8325 172.8108
Nevis Crossing CO -45.1756 168.9958
Ngatiawa River WN -40.9123 175.1230
Oaro KA -42.5164 173.5044
Oaro Beach KA -42.5094 173.5081
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TABLE 1. (Continued)
Locality (standard) AreaCode Latitude Longitude
Oaro, Omihi Stream mouth KA -42.4944 173.5169
Okuku River, 5km W of Loburn NC -43.2461 172.4665
Omoeroa River, S of Franz Josef WD -43.4028 170.0971
Oparara River NN -41.2000 172.1000
Opouri River, junction with Tunakino Valley Road SD -41.2000 173.6167
Otaki, 6 km S/SE WN -40.7911 175.1486
Otaki Forks WN -40.8333 175.2500
Otekaieke, Otekaieke River DN -44.8167 170.5667
Owhango, Whakapapa River TO -38.9959 175.3982
Palmerston, Shag River DN -45.4667 170.7167
Papatea, Waiti Stream BP -37.6649 177.8417
Pariwhakaoho River mouth NN -40.7740 172.7393
Peak 2.5 km W of Mount Tyndall OL -44.5310 168.6179
Pearce River Resurgence NN -41.2106 172.7403
Pelorus Bridge Scenic Reserve NN -41.2994 173.5733
Pelorus River, Daltons Bridge MB -41.2667 173.6333
Porters Pass, Lake Lyndon MC -43.3033 171.7025
Porters Pass, Lake Lyndon MC -43.2955 171.7111
Punakaiki, Bullock Creek BR -42.1014 171.3650
Rakaia River MC -43.7397 172.0306
Rotokohu, Inangahua River BR -41.9667 171.8667
South Highway 72, W of Staveley MC -43.6524 171.4455
Tangoio, near HB -39.3289 176.9125
Taupo TO -38.6861 176.0708
Te Awha Stream, Rangitukia Road GB -37.7347 178.4975
Temuka, Opihi River near Waiapi SC -44.2606 171.2289
The Remarkables, Rastus Burn CO -45.0553 168.8139
Thomas River, N branch MC -43.2086 171.7053
Totara River, near Moorhouse Road WD -45.9000 168.3833
Tukituki River, Pourerere Road, 10km E of Waipawa HB -39.9500 176.6833
Urewera National Park, Aniwaniwa GB -38.7453 177.1633
Urewera National Park, Te Taita a Makaro GB -38.6828 177.0569
Urewera National Park, Waimana River Valley, Pohatu Track, 3.2 km
E Tauwhare
BP -38.2911 177.1004
Vinegar Hill, Junction of Vinegar Hill Road & Rangitikei River RI -39.9167 175.6333
Waimakariri Gorge MC -43.3600 172.0503
Waimakariri Gorge, N of Waddington MC -43.3500 172.0500
Waimakariri River MC -43.4039 172.1100
Waioeka Gorge, N of Oponae and S of Opotiki BP -38.2005 177.3008
Wairau River, 2.5 km N of Leatham Road MB -41.6333 173.2167
Wairau River, Church Lane end MB -41.5333 173.5500
Wakarara, upper Waipawa River HB -39.7831 176.2656
Whakapapanui Stream, Junction Mahuia Rapid & Highway 47 TO -39.1333 175.5000
Whangaehu River, 5km N of Mangamahu WI -39.8000 175.3500
Winton, Oreti River SL -46.1333 168.2833
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Acknowledgments
For the opportunity to examine material in their care, the authors thank JW Early and D Ranatunga (Auckland
Institute and Museum, Auckland), PM Johns (Canterbury Museum, Christchurch), JMW Marris (Entomology
Research Museum, Lincoln University, Lincoln), RL Palma and P Sirvid (Museum of New Zealand Te Papa
Tongarewa, Wellington). Thanks are also extended to BE Rhode (Landcare Research, Auckland) for assistance
with digital photography and specimen databasing. This research was supported by Core funding for Crown
Research Institutes from the Ministry of Business, Innovation and Employment’s Science and Innovation Group,
for the Characterising Land Biota research portfolio.
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