The Nemesia trapdoor spider fauna of the Maltese archipelago, with the description of two new species (Araneae, Mygalomorphae, Nemesiidae)

. Contrary to what its name suggests, Nemesia arboricola is not strictly arboreal in habit. Here we compare female specimens of N. arboricola collected from arboreal and terrestrial nests. We furthermore describe the male of N. arboricola for the first time as well as two recently discovered species of Nemesia ( N. maltensis sp. nov. and N. cominensis sp. nov.). Nemesia maltensis is described from both sexes, N. cominensis is described from the female and a juvenile male specimen. For N. cominensis we discuss the sexual dimorphism of juvenile male and female spiders. Field observations and laboratory observations show remarkable features of the Maltese Nemesia fauna that are unknown from Nemesia species found elsewhere. Particularly, the arboreal dwellings of N. arboricola and the absence of a trapdoor to close off the burrow entrance in N. maltensis appear to be exceptional. The composition of the Maltese Nemesia fauna, as located in the central Mediterranean, is finally discussed in relation to the different Nemesia species-complexes occurring in the eastern and western Mediterranean basin.


Introduction
, is a Mediterranean genus of largely sedentary, terrestrial burrowing mygalomorph spiders that spend their lives in self-constructed, well camouflaged underground burrows (tunnels/nests) that are closed at the soil surface by a hinged 'trapdoor'. One longstanding and notable exception to terrestrial burrowing is Nemesia arboricola Pocock, 1903 from Malta. The original description of N. arboricola is based on a single female specimen (holotype in the Natural History Museum, London, see Kritscher 1994) that was reportedly found to construct its nest on the trunk of a tree (Pocock 1903). Over time the supposedly special arboreal habit of N. arboricola caused some confusion because Pocock had mistakenly imagined the nest of N. arboricola to be similar to arboreal nests that he knew from unrelated trapdoor spider species in the families Halonoproctidae Pocock, 1901, Migidae Simon, 1889and Barychelidae Simon, 1889(see Pocock 1897. In fact, these species construct very different tree-nests from that of N. arboricola. The main difference is that the species that Pocock envisioned do not excavate burrows, as all other trapdoor spider species including N. arboricola do, but construct short cocoon-shaped, or cigar-shaped nests of silk and debris that they attach to the surfaces of tree trunks or other hard surfaces (Griswold 1987;Schwendinger 2003;Decae et al. 2021). Such spider nests have never been found in Malta. Terrestrial burrowing Nemesia species however were long found to be common in the Maltese Archipelago, but because of their non-arboreal habits, the inhabitant spiders were not regarded to be N. arboricola. Instead, they were mostly taken to be N. macrocephala Ausserer, 1871or N. caementaria (Latreille, 1799 (Baldacchino et al. 1993). Fieldwork carried out in the last decennium of the 20 th century (Kritscher 1994) and the early 21 st century (Dandria 2001) however revealed that N. arboricola is not strictly arboreal and that the species excavates its burrows in various natural settings  like crevices between rocks and roots and, indeed, in soil or debris filled pre-existing holes in trunks of trees such as Phoenix dactylifera L., P. canariensis H.Wildpret, Olea europaea L. and Ceratonia siliqua L. These excavated 'tree-nests' of N. arboricola are qualitatively different from the nests of the Migidae, Halonoproctidae and Barychelidae mentioned above and from the nest that Pocock (1903) described in detail in his original paper on N. arboricola.
Here N. arboricola females collected from both arboreal and terrestrial nests are compared and their conspecific status is discussed. The male of N. arboricola is described for the first time. However, Baldacchino et al. (1993: fig. 1a), showing the distal palp and palpal organ of a male spider labelled "Nemesia (?) macrocephala" almost certainly is the first graphic representation of the N. arboricola male in literature.
We finally describe two newly discovered species of Nemesia. The first of these species is described from both sexes, while for the second species the adult male remains unknown, and the female and a juvenile male are here described.

Material and methods
Nest burrows of N. arboricola were located by visually searching the soil surface, tree trunks and rubble walls for the circular outline of their trapdoors. Females and juveniles of the two newly described species below were located by digging in soil with a small hoe; specimens of these species were taken alive and reared in plastic flower-pots filled with moistened soil in order to make observations of their burrows. For all three species, observations were made on the habitat, dimensions and internal structure of their nest burrows. Males were collected by hand when encountered in the field and by placing pitfall traps in locations inhabited by females. Retained specimens were stored in 70% ethanol. The study presented here is based on a sample of 21 Nemesia specimens collected between 1975 and 2021. For descriptive work, preserved individual specimens were placed in separate vials and given an individual identification number. TD-numbers for 3 ♀♀ and 1 ♂ collected in the 1970s and TC-numbers for 8 ♀♀, 8 ♂♂ and 1 juv. ♂ were provided accordingly (see sections Material examined below for individual numbers).
Microscopic observations were done with the aid of a Huvitz HSZ-645TR stereo microscope equipped with a Lusis HC-20CU camera operating on Panasis software and a Euromex iScoop compact microscope equipped with a Euromex VC-3031 camera. Both systems allow multiple focus photography and precision measurement. Habitus photographs of preserved specimens were made with an Olympus E-M5II camera equipped with an M. Zuiko Digital ED 60 mm F2.8 macro lens and a ring-flash. Figures were composed using Photoshop Elements 2021, lettering was done with Apple Preview. Observation techniques on preserved specimens are described in Decae et al. (2021).
Terminology follows Decae et al. (2021), except for terms that are of particular descriptive value for the genus Nemesia. Nemesia males carry distinctive 'armature' on the pedipalps and first legs that supposedly have a function in copulation and qualify as secondary sexual characters. These characters vary between species or species groups and therefore are valuable for intra-generic diagnostics. The following terminology and abbreviations are used to describe these characters here: -tibial spur (TS), a large, curved hook/spur, distally and prolateral on tibia I (Figs 29, 57) -tibial apophyse (TA), apical outgrowth of ventro-prolateral tibia I that carries the tibial spur (Figs 29, 57) -clasper field (CF), ventral side of curved proximal metatarsus I, furnished with short, stiff or spiky hairs (Figs 29, 57) -palpal tibial rake (PTR), dorsal apical group of strong distally pointing spines on the palp tibia 56).
Further morphological abbreviations used are: ALE = anterior lateral eye AME = anterior median eye AR = width anterior eye-row ATC = auxiliary tarsal claw av.
= compare with CL = carapace length CP = length cephalic part CW = maximum carapace width dia. = diameter dis. = distance EL = length of eye-group EmL = embolus length LL = labium length LW = labium width no.

New diagnosis
Female Nemesia arboricola from Malta is difficult to distinguish from N. macrocephala from nearby Sicily (Kritscher 1994). Females of the two species are among the largest Nemesia species known (fully grown females in both species range from 26 to 28 mm in total body length; the average in Nemesia is around 17.5 mm), and show only slight differences in their sexual and somatic morphology. Kritscher (1994) used the following three diagnostic characters to distinguish females of N. arboricola from those of N. macrocephala: (1) a different distance between the ALE and PLE, (2) a different shape of the spermathecal receptacles, (3) the presence or absence of labial cuspules. Pocock (1903) also reports the configuration of the eyes and the presence of labial cuspules as distinctive for N. arboricola. We found that, in general, the configuration of eyes and in particular the distance between the ALE and PLE (Figs 1-2 and 5-6) are unsuitable to distinguish N. arboricola from N. macrocephala on grounds of overlapping and highly variable measurements (dis. sd. 0.13 and av. 0.52,sd. 0.18,respectively). Although the shape of the receptacles appears to differ between the two species (Figs 3-4 and 7-8) it is difficult to quantify these differences. We therefore regard this character as only marginally reliable as diagnostic. We confirm Kritscher's third diagnostic character (also noted in Pocock 1903) as truly diagnostic and agree that the presence of labial cuspules in females is distinctive for N. arboricola . Furthermore, we found the dark coloured and speckled opisthosoma in N. arboricola  versus the lighter coloured opisthosoma with chevron stripes in N. macrocephala , and the thicker, slightly swollen, PMS  in N. arboricola as diagnostic characters to distinguish females of the two species.

Male
Because the male of N. macrocephala remains unknown, no characters that might distinguish this species from N. arboricola can presently be given. The male of N. arboricola however differs from nearly all Nemesia species, for which the relevant information is available, by the absence of the PTR (see section 'terminology' above). The only other species known in which the PTR is missing is N. simoni O. Pickard-Cambridge, 1874 from southwestern France and northern Spain. To illustrate the presence and/or absence of PTR in different species of Nemesia species, Figs 21-24 show the dorso-distal male palps of four different Figs 1-8. Characters reported to distinguish N. arboricola Pocock, 1903from N. macrocephala Ausserer, 1871in Kritscher (1994. 1-4. N. arboricola; represented by two specimens (TC.016, NHMR, Figs 1 & 3, specimen collected from an arboreal-nest; TC.017, NHMR, Figs 2 & 4, specimen collected from a terrestrial nest). 5-8. N. macrocephala; represented by two specimens (Isaia.046,Figs 5 & 7 and Isaia.047,Figs 6 & 8), both collected from terrestrial nests near the type locality, Palermo, Sicily. 1-2, 5-6. Comparative eye-formations. Note the overlap and variation in distances between ALE and PLE (ALE-PLE N. arboricola ranges between 0.10-0.20, n = 7; N. macrocephala ranges between 0.14-0.22, n = 2). Furthermore, note the variation in general configuration of the eyes that renders species recognition on these characters uncertain. 3-4, 7-8. Comparative shapes of spermathecae in ventral view. Note both similarities and differences in shape and differences in distance between receptacles of both species that renders unambiguous species recognition difficult, although spermathecal receptacles in N. arboricola shown here are somewhat smaller and finer built. Scale bars = 1 mm.
Nemesia species collected from geographically widely separated locations in the Nemesia distribution range (N. bacelarae Decae, Cardoso & Selden, 2007 from Portugal (Fig. 21), N. cellicola Audouin, 1826 from the Middle East (Fig. 22), N. simoni from southern France (Fig. 23) and N. arboricola (Fig. 24) from Malta). Note that species from opposite far ends of the distribution range and a new species from centrally located Malta (Fig. 56) feature a pronounced PTR only absent in N. arboricola and N. simoni . Nemesia arboricola males can be distinguished from those of N. simoni by their generally larger size (TBL » 13, n = 2 vs TBL » 11, n = 7) and on several aspects of its sexual and somatic morphology, its burrow structure and its distribution (compare the male description below with information given on N. simoni in Moggridge 1873). Figs 25-26 show distinctive differences in male palps (shape of tibia) and palpal organs of N. arboricola and N. simoni.

Holotype
In collection BMNH, London, not examined. See Kritscher (1994) for a detailed description and photographs of the specimen (in very poor and fragile condition), and for conformation that N. arboricola is the common and widely distributed cork-door building Nemesia species found in the Maltese Archipelago.  Pocock, 1903. 21-22. Dorsal palp tibia, PTR is present in all species of Nemesia for which information is available, here illustrated by two species from the eastern and western ends of the Mediterranean distribution range of the genus; N. cellicola Audouin, 1826 (21) from the Middle East and N. bacelarae Decae, Cardoso & Selden, 2007 (22) from Portugal. 23-24. Dorsal palp tibia of the only Nemesia species known in which the PTR is absent; N. arboricola Pocock, 1903 (23)   General coloration. Carapace uniform dark brown, chelicerae slightly darker than carapace, abdomen dorsal dark grey with light grey speckles in two parallel rows in the cardiac region ( Fig. 27), abdomen ventral light yellowish brown, palps uniform dark brown, legs dark brown proximally grading to light brown distally, sternum uniform yellowish brown, labium dark brown.
caraPace. Longer than wide (CW/CL 0.8), with light grey pubescence cover (partly lost), bristles concentrated along the margins and posterior on the thoracic part, cephalic part slightly elevated, thoracic part bulging slightly up from the fovea before sloping down to the posterior margin (Fig. 28).
chelicerae. Rastellum with few very strong apical teeth, teeth of diminishing strength and size along distal prolateral margin of chelicerae, single, prolateral row of furrow teeth, fang proximally bent, ventral serrated ridge.
oPisthosoma. Ovoid, anterior narrowing, covered with bristles, PMS small, close together and less swollen than in females, PLS short, thick, proximal segment longer than medial + distal segment, spigots restricted to apical spigot field.

Notes on female
Females of N. arboricola have been described in proper detail by Pocock (1903) and Kritscher (1994).
Here we compare specimens collected from tree-nests with specimens collected from terrestrial burrows. Figures 36-42, illustrate the variation in general appearance of the specimens in our study sample. Although individual variation in size, colour patterns and shades are evident, no qualitative morphological differences were found that would distinguish tree dwelling spiders from ground dwelling spiders at the species level. Moreover, all specimens in our sample closely fit the descriptions given by Pocock (1903) and Kritscher (1994). On these grounds we feel confident to state that arboreal and terrestrial specimens studied here are conspecific members of N. arboricola Pocock (1903). Figures 43-44, show characters common to all N. arboricola specimens (terrestrial and arboreal) studied, that distinguish N. arboricola from its supposed sister species, N. macrocephala.

Field observations
Nemesia arboricola constructs a simple tube-shaped burrow, capped with a sturdy lid (trapdoor) fitting neatly into the shaft (Figs 46, 51). Apparently older nests have been found with a layer of moss growing on the door surface (Fig. 45). The burrows are constructed in natural settings such as in tree trunks, between rocks in rubble walls and sloping ground (Figs 45-53). Tree-nests have been found between 150 cm and 8 cm above the ground surface in the trunks of ornamental palms (Phoenix canariensis and P. dactylifera), carob trees (Ceratonia siliqua) and olive trees (Olea europaea). In such tree-nests, spiders make use of pre-existing hollows and holes in the tree trunks into which they can snugly fit, or make use of hollows which are filled with woody debris and soil into which they can easily burrow. Burrows have also been found, at similar heights above the ground surface, in old rubble walls which hold back soil at field margins; here the spiders burrow into the clayey soil which fills the cracks between the individual stones. In both tree-nests and wall-nests, the burrow lids are flush with the vertical surface and dorsally hinged so that the trapdoor opens vertically upwards, with the shaft directed into the trunk/ wall perpendicular to the vertical axis. Nemesia arboricola also nests directly in the ground, but always it seems under one of the following conditions: the ground must be steeply sloping or, if the soil is flat, located immediately beneath an overhanging structure such as large rocks or a rubble wall (Figs 52-53). No burrows have been found on level ground in full exposure ('in plain sight' as it were). Tree-nests and wall-nests vary from 4.5 cm to 9.5 cm in depth for mature individuals, with burrow width varying according to the size (maturity) of the individual. Ground-nests in deep soil have, on average, deeper shafts than tree or wall-nests. In all cases, the burrow shaft is relatively simple in construction; it has entire, thick lining of silk with no internal doors, plugs or other features; and it is unbranched.
The resident spiders themselves respond to disturbance by holding the hinged door very tightly; if the door is prised open using forceps (which takes considerable force) the spider may either retreat immediately head-up to the bottom of the shaft, or else repeatedly attempt to close the door while striking the forceps aggressively with its chelicerae. Recently hatched juveniles are retained within the mother's burrow, from which they disperse and apparently establish themselves in close proximity, as N. arboricola populations are always localized and often quite dense. Moulted exoskeletons are discarded from the burrow and may be observed outside near the burrows if they are sheltered from wind. During the aestivation period, the door to the burrow is sealed shut from the inside, but the door still remains exposed as under normal circumstances (i.e., the burrow lid is not concealed with soil during periods of inactivity). Most mature males emerge from their burrows and wander between the months of November and March, though a few individuals may appear outside this timeframe.

Etymology
The name refers to Malta, the largest island in the Maltese Archipelago and the only Mediterranean island where the species is currently known to occur.
General coloration. Carapace yellowish with dark flanks of cephalic part and dark margin (Fig. 55), chelicerae brown, darker than carapace with dorsal lighter coloured patch, opisthosoma dorsal anterior dark greyish-brown, further light coloured with dark, irregular chevrons and a dark cardiac line (Fig. 54), opisthosoma ventral uniform creamy white, palps and legs lighter coloured than body in dorsal view and ventrally lighter coloured than dorsally, sternum and ventral coxae creamy white, labium conspicuously darker than other ventral parts.
caraPace. Longer than wide (CW/CL 0.8), covered with very fine silvery pubescence, curved bristles concentrated along the margins and posterior on the thoracic part, longitudinal row on the crest of the cephalic part and few forwardly projecting bristles on clypeus. Cephalic not elevated, thoracic part bulging slightly up from the fovea before sloping down to the posterior margin (Fig. 55).
chelicerae. Rastellum with few strong teeth on the apical prolateral corner of chelicerae, prolateral row of furrow teeth, fangs sharp, bent, ventral un-serrated ridge.
General coloration. Carapace cephalic part has wide light brown crest-zone and is generally darker in colour than thoracic part, black margin prominent (Fig. 64), chelicerae bicoloured brown, slightly darker than carapace, abdomen dorsal as in male, palps and legs generally as in male, but dorsal femora lighter coloured and with prominent maculae (Figs 64-65, 67), sternum and labium slightly darker coloured than ventral coxae.
caraPace. Longer than wide (CW/CL 0.7), sparsely covered with very fine black pubescence, bristles centrally in longitudinal row on crest-zone. Cephalic part only slightly elevated and gradually sloping down over the thoracic part to the posterior margin (Fig. 65).
PalPs. Femur and patella spineless, tibia ventral and prolateral sharp distally pointing spines and dorsal long parallel rows of trichobothria, tarsus, distally pointed, ventral half fully scopulate with group of sharp distally pointing spines, dorsal row trichobothria in central part of article, palpal claw with short proximal row of teeth.

Observations
In the field (Fig. 70), the burrows of this species appeared to be completely lidless, with the burrow entrance opening at the ground surface directly (Fig. 72). The wall of the burrow is lined with almost imperceptibly thin strands of silk from the inside; but the burrow walls derive most of their structural integrity from the compaction of the soil around them (Fig. 71). In captivity, once soil was moistened, a female constructed a bifurcated burrow with two shafts opening at the soil surface with no lids (Fig. 73), joining together into one shaft deeper into the soil. Females were collected from burrows some 15 cm deep in the soil, congregated at the edge of a small boulder embedded in deep soil in a wooded area populated by trees of Laurus nobilis L., Olea europaea, Rhamnus alaternus L. and Pinus halepensis Mill. Burrow entrances always appeared at the soil surface at the edge of the boulder or, in one instance, at the base of the thick stems of an Acanthus mollis L. plant nearby. Males were collected in pitfall traps after several heavy rainfall events which occurred in September -the so called "first rains" which bring an end to the dry season.

Diagnosis
Nemesia cominensis females can be distinguished from all known Nemesia species by its general light colour (Figs 74-75) and the elongated, wiggly, tube-shaped spermathecal receptacles without a clear differentiation in proximal, medial and distal parts (Figs 79,87). It further differs from N. maltensis in the absence of maculae on legs (Fig. 75) and the elevated cephalic part (Fig. 75).

Etymology
The name refers to Comino, the third largest island in the Maltese Archipelago and the only Mediterranean island where the species is currently known to occur.

Description
Female holotype (no. TC-002, NHMR) General coloration. General appearance as a relatively light-coloured Nemesia species (Fig. 74). carapace cephalic part with wide orange-brown crest zone and light grey flanks, thoracic part light yellow with vague grey folia pattern, chelicerae dark brown, distally darkest, palps and legs crème-coloured with dorsal yellow zones, sternum light yellow, labium and maxillae light brown, opisthosoma dorsal creme-colour with light grey pattern of blotches and chevrons, ventral orange-brown zone between epigastric furrow and light-coloured spinnerets.
caraPace. Longer than wide (CW/CL 0.8), sparsely covered with very fine black pubescence, bristles centrally in longitudinal row on crest-zone and around the eyes. Cephalic part elevated; fovea only weakly recurved. Eyes: eye-group almost twice as wide as long (EL/PR 0.48), PR slightly wider than AR (PR/AR 1.03), AME slightly more than their diameter apart (dis.AME/dia.AME 1.10), distance ALE-PLE less than ½ dia.ALE (ALE-PLE/ALE 0.44).
PalPs. Femur and patella spineless, tibia two ventro-lateral rows of sharp distally pointing spines and dorsal long parallel rows of trichobothria, tarsus ventral half fully scopulate with group of sharp distally pointing spines, dorsal trichobothria in V-formation, palpal claw with short proximal row of four teeth.
Sub-adult male (no. TC-003, NHMR) As in all species of Nemesia, sub-adult spiders are, except for their small size, similar to adult females in general appearance and somatic characters. Males and females of Nemesia can be distinguished in juveniles by internally checking the presence of spermathecae that are already detectable in small juvenile and sub-adult females (Fig. 87 cf. Fig. 88). Sub-adult males overlap in general body size with young females, but can be distinguished on the shape of the palp-tarsus that is slightly swollen (Figs 82,84 cf. 81,83) and the absence of a vulva structure in the epigastric furrow (Fig. 85

Observations
Nemesia cominensis constructs a thin, flimsy wafer-door lid to its burrow (Fig. 90), which is partly concealed by a thin layer of loose soil particles on top (Fig. 89). Nemesia cominensis captures its prey in the usual trapdoor spider fashion by laying in ambush under a cracked open trapdoor, launching a flash attack on small animals that wander within reach of the spider (Figs 91-94). Burrows are constructed in the ground in deep soil, with the door flush with the flat ground surface, and may be unbranched (single shaft) or bifurcated (two shafts joining to become one at the bottom, in a Y shape). During the aestivation period, the burrows have no lids whatsoever, and the burrow entrances are instead concealed and obstructed by up to about 2 cm of soil; they are also completely devoid of any silk lining. When the soil is moistened, the spiders resume their door-making and hunting; the first few millimetres of the burrow entrance are lined with a very thin layer of silk to which the door is attached, but the rest of the burrow still remains without silk lining. It is not known how deep the burrow shafts are in nature during the active period; in captivity an active burrow was constructed which reached 8 cm in depth but this was restricted by the size of the flower-pot; on location some spiders were found residing in aestivation burrows up to some 15 cm deep into the soil. Recently hatched spiders have been found after digging up aestivation burrows belonging to mature individuals, so it is assumed that the young are retained within the maternal burrow and disperse afterwards, taking up residence in close proximity (the sampled population was localized and dense). So far, this species has only been found to construct its burrows in relatively deep soil at the bottom of a shallow valley on the island of Comino, from which all of the aforementioned observations have been made. The difficulty in locating specimens may doubtless obscure the true distribution and ecological preferences of this newly described species from the Maltese Islands.

Discussion
Within the genus Nemesia, different regionally distributed species-complexes and further sub-groups can be distinguished (Simon 1914;Decae et al. 2015). Differences between the Nemesia species compositions of the eastern and western basins of the Mediterranean are particularly distinct in this respect (Decae 2012). Located in the centre of the Mediterranean where the different eastern and western Nemesia faunae meet (Decae 2012), the Maltese Nemesia species appear to take an intermediate position between East and West. Judged on morphology, N. arboricola has its closest relatives in the western Mediterranean (Decae 2012 group CF), whereas N. maltensis sp. nov. and N. cominensis sp. nov. Are closer to eastern Mediterranean Nemesia species (Decae 2012 group AD). Malta (and adjacent regions in Sicily, Italy and Tunisia) may therefore act as focal point for future taxonomical, distributional and phylogenetic research aimed at elucidating the origin, intrageneric relations and dynamics of the highly diverse, complex, and yet understudied, Mediterranean Nemesia fauna.
Two other points that are worth discussing concern the apparently unique habits of N. maltensis sp. nov. and N. arboricola. Firstly, N. maltensis sp. nov. appears to be unique among Nemesia species in its nest building behaviour by constructing an underground nest without a trapdoor cover (Figs 71-73). Although open burrows have not been reported before for Nemesia species, such burrows are known from the related Mediterranean nemesiid genus Brachythele Ausserer, 1871. Brachythele and Nemesia have overlapping distributions in the north-eastern Mediterranean (personal observation, AED). Secondly, N. arboricola is the only arboreal Nemesia species currently known (Figs 47-51). As for the observed inclination of N. arboricola to construct its burrows in substrate-fillings of steep or vertical surfaces (rubble walls, tree trunks) it is worth noting that Balearic species such as N. bristowei Decae, 2005 andN. randa Decae, 2005 are also found to exploit vertical surfaces as burrow locations (Decae 2005). These species are commonly found in rubble walls on steep cliffs and even on overhanging surfaces (personal observation, AED) and might be expected to be found in trees as well, given the presence of suitable soil fillings.
A final point to be discussed is the information on the phenology of Nemesia species. The two males of N. arboricola (from our samples) were collected in different seasons (November and March respectively) indicating that there is no particular mating season for this species. A sample of 124 male collection date records, obtained from populations occurring throughout the Mediterranean shows that adult Nemesia males may be found all year round with the possible exception of midsummer. However, two waves of male collection/wandering are evident from our data. About one quarter (27%) of the males were collected in the first half of the year and about three quarters (73%) in the second half of the year with a peak in the months of September and October. Available data are currently insufficient to discern seasonal instances of male emergence at the species level.